all issues addressed!
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112
rebuttal2.tex
112
rebuttal2.tex
@@ -54,7 +54,8 @@
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like to offer several suggestions that may serve to either enhance
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the manuscript or inspire future research endeavors.}
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\response{}
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\response{Thank you for trying to make our manuscript more
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biologist-friendly!}
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\issue{First, I should point out that beyond the presence of a
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threshold-induced nonlinearity, the complex structure of the
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@@ -67,7 +68,7 @@
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in Cellular Neuroscience, 13, 354. for a discussion of the general
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case and the study by Troy Smith, Unguez and Weber (2006, Fig. 3) in
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which receptor cells of tuberous electroreceptor organs and their
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afferents from Apteronotus leptorhinchus were labeled to varying
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afferents from Apteronotus leptorhynchus were labeled to varying
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degrees by six anti-Kv1 antibodies. Kv1.1 and Kv1.4 immunoreactivity
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was intense in the afferent axons of electroreceptor organs. It is
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noteworthy that Kv1 are low-threshold channels and, in some cases,
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@@ -76,16 +77,15 @@
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strengthen the links between well-written theoretical analysis and
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the practical field of experimental physiology.}
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\response{READ PAPERS AND CITE THEM You are right, there are more
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nonlinear mechanisms potentially contributiong to the threshold
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nonlinearity. We now mention this in the methods when introducing
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the threshold nonlinearity (after eq. 13) and cite the corresponding
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manuscripts. We also added a paragraph to the methods reiterating
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what we have in the discusssion, that the details of this
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nonlinearity are not so important in the context of the present
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manuscript, since we compute all cross-spectra between the resulting
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amplitude modulation and the spikes responses. MAYBE ALSO IN THE
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CONCLUSION.}
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\response{You are right, there are more nonlinear mechanisms
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potentially contributing to the threshold nonlinearity. We now
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mention this in the methods when introducing the threshold
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nonlinearity (after eq. 13) and cite the corresponding
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manuscripts. We also added a sentence there reiterating what we have
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in the discussion, namely that the details of this nonlinearity are
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not important in the context of the present manuscript, since we
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compute all cross-spectra between the resulting amplitude modulation
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and the spikes responses.}
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\issue{Second, and along the same lines, the discussion could be
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improved by mentioning the effects and significance of these
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@@ -94,7 +94,8 @@
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interference avoidance responses, and b) transient changes, as in
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chirps.}
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\response{HERE: STATIONARY ANALYSIS, CHIRPS ARE TRANSIENT, NEEDS RESEARCH. JAR SIGNALS are slow but change continously so they might hit the baseline frequency Add something somewhere around line 660.}
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\response{We added a paragraph addressing JARs, chirps, and rises to
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the discussion (lines 695 -- 703).}
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\issue{Finally, the precise description of the methods could be
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expanded for reaching a broader biology audience; in particular, the
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@@ -103,8 +104,6 @@
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first reading of the methods, although accurate, does not offer the
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biology reader a quick and intuitive approach to the study.}
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\response{IMPROVE METHOD DESCRIPTION AS DESCRIBED BELOW}
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\issue{Next, I list some minor more detailed comments that may clarify
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the design and methods and facilitate their understanding by a
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broader audience.}
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@@ -118,11 +117,12 @@
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between the extreme profiles of P (signal amplitude) and T (signal
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slope)?}
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\response{In \textit{Apteronotus} T-units are characterized by 1:1
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locking to the EOD, i.e. by having a baseline firing rate matching
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the EOD frequency. We definitely have no T-units in our data
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set. This we explain now in the ``Identification of P-units and
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ampullary cells'' section in the methods.}
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\response{T-units are characterized by 1:1 locking to the EOD, i.e. by
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having a baseline firing rate matching the EOD frequency. We
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definitely have no T-units in our data set, since our P-unit firing
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rates are well below the EOD frequencies. This we explain now in the
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``Identification of P-units and ampullary cells'' section in the
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methods.}
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\issue{In line 147, rather than using the term
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"laterally," I believe it would enhance clarity to state "parallel
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@@ -137,7 +137,10 @@
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field foveal perioral region where the majority of receptors are
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located.}
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\response{ADD SOMETHING TO STIMULATION SECTION LINE 110}
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\response{As stated in ``Experimental subjects and procedures'', all
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recordings were done in the posterior lateral line nerve. So we did
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not record from the foveal perioral region, and hence this problem
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is not relevant.}
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\issue{Line 148, the phrase "band limited white noise" lacks
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clarity. Upon my initial reading, I assumed that the cutoff limit
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@@ -153,7 +156,8 @@
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could benefit from greater clarity to avoid the need to explore the
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results first in order to understand well.}
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\response{STATE TYPE OF FILTERING IN STIMULATION SECTION, CITE ALES SKORJANC}
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\response{We added a sentence that describes how we generate those
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stimuli in the Fourier domain (lines 156--160).}
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\issue{Line 154. This procedure elicits a modulation of the envelope
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of the reafferent signal. To achieve this, you adopted distinct
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@@ -163,7 +167,10 @@
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averaged sine wave recorded via local electrodes adjacent to the
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gills exhibited an increase of 1 to 5\%, is this correct?}
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\response{NO! EXPLAIN AND ENHANCE STIMULATION SECTION}
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\response{No! We increased the amplitude of the white noise until the
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standard deviation (not the mean) of the resulting modulation of the
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EOD reached 1 to 5\,\%. We rephrased the description of the
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stimulation and hope that this is clearer now (lines 164--168).}
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\issue{b) with regard to P receptors, you multiplied the head-to-tail
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ongoing signal by a white noise signal and played the resultant
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@@ -184,7 +191,17 @@
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on the peaks of the signal themselves? How does this affect the
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recruitment of P and T receptors?}
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\response{ALL THESE DETAILS DO NOT MATTER AT THE LEVL OF INDIVIDUAL P-UNITS. SEE HLADNIK.}
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\response{You are right about the phase shifts and that this does not
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``significantly impact individual receptors response''. This is a
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standard stimulation procedure for characterizing receptor responses
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that are located mainly on the sides of a fish's flat body. See, for
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example, Hladnik and Grewe, 2023. And yes, this will probably impact
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relative spike timing in distinct receptors and thus may also impact
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the JAR mechanisms. However, this manuscript is about single
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receptor responses and not about T-units, and we feel it is already
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complicated enough. Therefore we would rather prefer to not open up
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all these issues, since they are not relevant for the results we
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present.}
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\issue{Line 238. Are you referring to the terminal non-myelinated
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branches that connect receptor cells to the initial Ranvier node?
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@@ -200,7 +217,9 @@
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organ. Could you discuss this aspect, considering the anatomical
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structure of the receptor in your species?}
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\response{READ LITERATURE AND SAY A FEW WORDS}
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\response{Exactly. We slightly expanded our description to make clear
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that we talk about the signal transduction until it reaches the
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spike initiation zone (lines 258 -- 259).}
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\issue{\large Reviewer \#2}
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@@ -252,15 +271,16 @@
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clipped in these two figures.}
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\response{You are right. In figure 4 we show now the spectrum up to
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750\,Hz, such that $f_{EOD}$ and its interactions with $\Delta f_2$
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and harmonics are included. We labeled the additonal peaks
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accordingly. In figure 3 we stay with the small range, because we
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have so little data for this special setting where one of the beat
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frequencies approximately matches the P-units baseline firing rate
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(only three trials of 500ms duration). This is why the power
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spectra are very noisy. Also, for an introductory figure we prefer
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to only show the few peaks that are relevant for the rest of the
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manuscript, such that the reader does not get overwhelmed.}
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950\,Hz, such that $f_{EOD}$ and its interactions with $f_1$ and
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$f_2$ are included. We labeled the additional peaks and expanded the
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figure caption accordingly. In figure 3 we stay with the small
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range, because we have so little data for this special setting where
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one of the beat frequencies approximately matches the P-units
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baseline firing rate (only three trials of 500ms duration). This is
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why the power spectra are very noisy. Also, for an introductory
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figure we prefer to only show the few peaks that are relevant for
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the rest of the manuscript, such that the reader does not get
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overwhelmed.}
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\issue{(6) Figure 3. Why are these example firing rates based on
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convolution with a 1 ms Gaussian kernel if the analyses were based
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@@ -269,13 +289,13 @@
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actually analyzed. More fundamentally, why would a 2-fold difference
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in kernel width be appropriate for presentation vs. analysis?}
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\response{This was for historical reasons. We now decided to use the
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\response{This was for ``historical'' reasons. We now decided to use the
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1\,ms kernel for all figures and analysis. In doing so we also added
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panels showing firing rates in addition to the response spectra in
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figure 4. Using the more narrow kernel better reveals the details of
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the time course of the firing rate and this way improves the
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connection between the firing rate and the response spectra. In
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figure 10, middel column, the range of possible values of the
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figure 10, middle column, the range of possible values of the
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response modulations is a bit enlarged by using the 1\,ms kernel,
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but the correlations and their significance did not change a lot
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either.}
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@@ -289,12 +309,12 @@
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stimulus. We added a sentence to the end of the figure caption to
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make this clear.\\
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If it were the power spectrum of the signal after it passed
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a non-linearity (rectification or threhsolding at zero), then there
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a non-linearity (rectification or thresholding at zero), then there
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could be also peaks at the sum and difference of the beat
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frequencies. However, since they are close to the higher one of the
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two beat frequencies they do not show up in the AM as obviously as
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for the settings used in the social envelope papers by Eric Fortune
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and Andre Longtin and colleges (I guess this is what you have in
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and Andre Longtin and colleges (I guess this is what you had in
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mind).}
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\issue{(8) Line 302. "not-small amplitude" is arbitrary and
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@@ -324,12 +344,12 @@
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\response{Yes, in figure 4 increasing stimulus contrast results in
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stronger nonlinearities. There the stimuli are narrow-band sine
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waves. However, as pointed out in the context of figure 7, when
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using a broad-band noise stimulus instead, this stimulus by itselfs
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using a broad-band noise stimulus instead, this stimulus by itself
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adds background noise to the system that linearizes the
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response. That is why the susceptibilities estimated from noise
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stimuli decrease for higher stimulus contrasts.\\
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We added a whole paragraph at the beginning of this section to make
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this clear.}
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this clear (line 477 -- 482).}
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\issue{(12) Lines 655-675. This was a very nice end to the discussion,
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but I would like to see more. I would like the broader significance
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@@ -343,6 +363,16 @@
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reiterate these points briefly and delve into more detail on
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comparative considerations.}
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\response{UH. LETS THINK ABOUT IT.}
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\response{We also like to see more on this, but we feel that we
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already speculated enough. Without further studies on the readout of
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the receptor responses, we cannot make any convincing claim about
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whether and how weakly nonlinear interactions are actually utilized
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in a neural system. The problem is that a match of one of the
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stimulating frequencies or their sum with the neuron's baseline
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firing rate is required. This is all addressed in the (now second
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final) paragraph of the ``Nonlinear encoding in P-units'' section.
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However, we agree that the comparative aspect of the conclusion
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could be expanded. We therefore added one more final speculative
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sentence to the conclusion.}
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\end{document}
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