Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Celestial polarization patterns sufficient for Viking navigation with the naked eye: detectability of Haidinger’s brushes on the sky versus meteorological conditions Gábor Horváth, Péter Takács, Balázs Kretzer, Szilvia Szilasi, Dénes Száz, Alexandra Farkas and András Barta Article citation details R. Soc. open sci. 4: 160688. http://dx.doi.org/10.1098/rsos.160688 Review timeline Original submission: Revised submission: Final acceptance: 10 September 2016 9 January 2017 9 January 2017 Note: Reports are unedited and appear as submitted by the referee. The review history appears in chronological order. Review History RSOS-160688.R0 (Original submission) Review form: Reviewer 1 (Shelby Temple) Is the manuscript scientifically sound in its present form? Yes Are the interpretations and conclusions justified by the results? Yes Is the language acceptable? Yes Is it clear how to access all supporting data? Yes Do you have any ethical concerns with this paper? No © 2017 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 2 Have you any concerns about statistical analyses in this paper? No Recommendation? Accept with minor revision (please list in comments) Comments to the Author(s) Comments to Authors General Comments This is a great analysis of the proportion of the sky that would be useful for human navigation using celestial polarisation patterns relative to meteorological conditions. My only major concern with the manuscript is the comparison of the three different wavelength bands used. It is not necessary to provide all three colour bands, and in fact it detracts from the story by making it more complicated than it needs to be. The entoptics phenomenon of Haidinger's brushes is mediated by the absorption of short-wavelength light by the macular carotenoids (lutien, zeaxanthin and meso-zeaxanthin) (Müller et al 2016, and Bone 1984) and is not visible at wavelengths above about 500 nm, therefore the two colour bands in the green and red are not useful for perception of polarization and therefore do not add to your argument. Instead the extra data makes the story more confusing. If you remove the colour component you can focus on the main effect in the blue and make the story more concise and more easily accessible. In case you are interested, the original experiments we did (Temple et al. 2015) were done with humans looking at a modified LCD screen, however, I have since measured the human threshold using a sheet of rotating polarizer combined with a series of my custom made depolarizing filters and found that the human threshold is even lower than 23% we originally reported. Out of 20 people studied, four individuals could see HB down to 16% and one could go as low as 12%. In this study the mean d* was 32.4% and the min was 70%. If you want to add these results to this paper you could report them as unpublished data or personal communication. This work was done as part of a preliminary experiment that showed a correlation between threshold for detecting HB (d*) and macular pigment optical density (MPOD) measured on a dual wavelength scanning laser ophthalmoscope (Heidelberg Spectralis). These results concur with recent findings of Müller et al. 2016 (Müller PL, Müller S, Gliem M, et al. Perception of Haidinger brushes in macular disease depends on macular pigment density and visual acuity. Invest Ophthalmol Vis Sci. 2016;57:1448–1456. DOI:10.1167/ iovs.15-19004). It is not helpful for the reader when sentences have too many equations (e.g. P5 L5-11 ) or are written around a figure (e.g. start with "Figure x shows..."). It is much easier to read and understand the point being made if the sentences are written in words and just tell the story, leaving the equations and figures to the end to be put in parentheses e.g. Page 3 Line 56 "Figure 2 shows a totally overcast sky taken with our full-sky imaging polarimeter when the solar elevation was θ = 0º and the cloud percent was ρ = 100 %." It is much easier to understand if you focus on the results not the figure, as I have tried to do here "When skies are clear the proportion of the celestial hemisphere P suitable for human polarimetric navigation based on perception of Haidinger's brushes is high (insert P-value here; Figure 2). However, when the sky is overcast, P approaches 0 making it infeasible for navigation by this means (Fig. 3)." NOTE I would reverse the order in which these two points are presented and start with the positive situation in which polarimetric navigation IS possible rather than starting with when it IS NOT. Specific Comments To preclude, I really like this work and think that it will be a valuable contribution to the field but that it needs to be written in a way that makes it more accessible to the reader, so I have provided numerous suggestions below. Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 3 Page 1 Title The title is long and not easily understandable, specifically the expression "sky-polarization prerequisite" is not clear. Please consider an alternative, for example a variation of: Limits of human polarization perception support Viking navigation with the naked eye based on celestial polarization patterns. OR Celestial polarization patterns sufficient for Viking Navigation with Haidinger's brushes. OR Viking navigation with Haidinger's brushes: assessment of the meteorological conditions and proportion of the sky suitable based on human threshold for polarization perception. Page 1 Line 25 (P1 L25) Here 'brush' is used rather than "brushes" I recommend brushes but pick one and stick to it for consistency. P1 L25 you have already said '8-shaped' and I am not sure that 'brush-like' is a good description, I recall that the word brush was a poor translation of the German that was meant to be bushel, like a bushel of wheat. A better description is 'bowtie-like' rather than 'brush-like' P1 L25-28 This Sentence is long and difficult to understand with novel terminology that needs defining upon first use. "Since the celestial great circle passing through any sky point with high enough d-values and parallel to the yellow Haidinger’s brush crosses the sun, if two such sky points are selected, the sun position can be determined, which is at the intersection of the two great circles." specifically you need to define or come up with a better description of "the celestial great circle" And this statement is really unclear "passing through any sky point with high enough d-values and parallel to the yellow Haidinger’s brush crosses the sun" please see if you can reword this. The last part refers to "two great circles" but the first part makes it seem like there should only be one "the celestial great circle". Personally, I can see HB quite easily both in clear skies and in the forest when only a very small portion of the clear sky is visible through the canopy, and I don't think it is necessary to talk about 'great circles' but rather I think you could talk about a band of high degree of polarization that arcs across the sky at about 90 degrees to the position of the sun and that a person could pick two points on that band and observe the yellow brushes of HB and from that triangulate the position of the sun based on the orientation of those two observed brushes. But equally it does not need to be static observation. by sweeping your eye across the band of high DoP it is possible to get a dynamic view of HB that point directly at the sun. P1 L30 in addition to occlusion by cloud and fog, HB can also be used to locate the sun when it is below the horizon or occluded by objects on the horizon. P1 L30 the expression 'atmospheric optical prerequisite' is overly complicated, and this sentence could be simplified to something like "In order to determine the position of the sun using the celestial polarization pattern the d of the portion of the sky used, must be greater than the viewers degree of polarization threshold (d*) for perception of Haidinger's brushes." Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 4 P1 L34 remove the red and green here and throughout the paper (see general comment above) P1 L36 recommend changing 'cloud percent' to 'cloud cover (in percentage)' P1 L37 the statement 'We obtained that P is large enough only for the most sensitive human observers having perception threshold d*min = 23 % of Haidinger's brushes." does not make sense. "Large enough" for what? P1 L38 no space required between the number and the percent symbol, whereas there is a space required between a number and most of other units (e.g. nm) but not degree. P1 L39 recommend change to 'with cloud coverage of p = 0%, when the sun's position is already easily determined." delete "since it is visible with a big chance". P1 L42 again what does 'P is large enough' mean? if P is the proportion of the sky where d>d* then surely an observer only needs a couple of small areas of the sky that are larger the 5 degrees in radius (size of HB perception) to observe HB and detect the position of the sun, so if the d is high then only a small portion of the sky located near the band of maximum d is required. So what does 'large enough' mean? P1 L45 change 'great' to 'large' P1 L48 because on-line search engines will search your title and abstract it is redundant to use keywords that are already in your title and abstract in your list of keywords. Instead consider using other words that a potential reader might use to search for similar articles. e.g. calcite, orientation, sea-faring, orienteering, migration, vision, macula, macular pigment etc... P1 L53 see comment above about 'brush-like' could add that the two 8-shaped figures combine to form a Maltese cross shape P1 L54 & L55 stick to the '8-shaped figure' or perhaps 'bowtie-like shape' as used elsewhere P1 L55 delete 'well-known'. Haidinger's brushes are certainly not well known. I have been interested in this subject for near 20 years and very few vision scientists, optometrist, visual ecologist have ever heard of them and even few of the general population. P2 L2 change to 'brushes' here and throughout. P2 L2 suggest removing the word 'well' most people find it difficult to perceive HB at the best of times, and even those of us that can see them consistently would not say they are easy to see or are highly salient, it suffices to say that they can be perceived. P2 L4 suggest change to "when large portions of the sky are red and orange, or when the sky is..." P2 L11 correct 'treshold' to 'threshold' P2 L10 it is much easier to read if words are used rather than equations e.g. 'Recently, Temple et al [13] measured the degree of polarization threshold, d*, at which humans could just detect Haidinger's brushes. P2 L14 consider adding to this sentence as such "determined by the relative position of the observer, the sun and the celestial point observed (Fig. 1)." P2 L21 this 'celestial great circle' needs to be well defined earlier in the paper if it is going to be used. Otherwise consider 'band of highest polarization' Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 5 P2 L24 if is not clear why there are now 'two great circles'. I know what you are talking about but it is not written clearly enough for someone slightly unfamiliar with the specifics to understand it well enough. Furthermore there are no 'great circles' in your figure 1, nor should there be. I think it easier to describe it somewhat as I have in my comment above re P1 L25-28. P2 L34 change to 'they had their own analyser..." P2 L43 change to past tense 'the aim of this work was to study' P2 L44-45 remove red and green from here and the paper in general P2 L47 change to 'percent cloud cover' here and throughout the manuscript P2 L50 used past conditional past tense ' navigation could have been based on the perception...' P3 L14 change to 'coverage of the visible sky estimated by visual examination [40].' P3 L15 suggest 'refer to ever increasing cloud coverage' P3 L24 spelling 'approximately' P3 L30-32 this is a bit confusing, it is clear that there were 12 sun azimuth groups and 9 cloud coverage groups, but what are the '12 different skies"? Are these just 12 examples/samples of each of the above 108 groups? if so then this needs to me spelled out (described) more clearly. P3 L32-34 delete reference to colour P3 L43 consider 'probability that a viewer (Viking) with high, average and low sensitivity, respectively, ...' P3 L45 remove reference to colour P3 L50 the statement says that the areas that were not sky were masked out but they still show up in the figures (e.g. Fig 2 c,d, & e and 4 c,d & e) with red and blue regions suggesting under and overexposed regions, but really this just detracts from the message you are trying to get accross, so I recommend blacking out the non-sky regions in these images to remove the red a blue that is not in the sky (the only part of interest here). P3L57 see note in general comments about the order of presentation of results. Start with the positive situation i.e. clear skies (section 3.2 rather than 3.1) and use full sentences that do not start with "Figure X shows..." Actually by the time you remove the parts of 3.1 and 3.2 that belong in discussion and remove the reference to red and green the sections will be short enough that they can be combined into a single section that just focuses on percent cloud cover. P4 L9-12 this statement is interpretation of the results and therefore belongs in discussion section not results. P4 L16 change sentence so it does not start with "Figure 4A..." P4 L22-29 this statement is interpretation of the results and therefore belongs in discussion section not results. P4 L31 consider changing to "sun elevation-cloud cover matrix..." Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 6 P4 L33 change sentence so it does not start with "Figure 6..." P4 L44 change sentence so it says something more like "A person having low polarization sensitivity (e.g. d*max = 87%) can only detect HB under a select few meteorological conditions (Fig 6; Supplementary table S3, S6, S9) and would therefore be an unlikely choice for a Viking navigator." In the discussion, you could suggest that Viking navigators were likely those that had the highest polarization sensitivity (So move P4 L59 to discussion), you could also add that a diet rich in fish, which is a good source of macular carotenoids, may have contributed to Viking's general ability to see HB. P4 L50 same as above. P5L5-11 this paragraph is almost incomprehensible, it is written as an equation masquerading as a sentence. Rewrite as a sentence with words rather than symbols. P5 L20 This paragraph starts off well with clear words used to describe everything and then the reader gets to this line that starts with <P>(θ,ρ), and gets totally lost. Please write out in words what these symbols mean as you did in the first two sentences. P5 L25 need to change this to blue since polarization sensitivity has nothing to do with the human spectral sensitivity curve that peaks in green, and everything to do with the polarization sensitivity, which is maximal at 460 nm (2nd sentence of Chapter 14 of your book Gábor that we wrote together) due to absorption of macular pigments (Bone 1980 Vision research 20:213-220). The statement that blue can be ignored is incorrect. In blue light the HB become clearer, and it is thought that the yellow colour a person sees under full spectrum illumination is a psychophysical effect of the eye where yellow is perceived when there is an absence of blue. I agree that the blue brushes of HB are or little utility for navigation but that has nothing to do with the spectral content of the sky. P5 L31 & 34 again consider rewording the 'great circle passing the sun', especially the last part 'passing the sun' this does not make sense. P5 L35-45 suggest changing to something like... "We found that the proportion of the sky suitable for the humans with a high sensitivity to Haidinger's brushes (d*min = 23%) is high under numerous meteorological conditions, particularly when the sun is low on the horizon and when the sky is clear of cloud. When the sun is below the horizon, and therefore not possible to see or accurately locate, the proportion of the sky suitable for sky polarimetric navigation is between 40% and 77% depending on cloud cover. P5 L48 Need to define "3rd step" of just use the word-based description first and then put '3rd step' in brackets and state that this is what they called it. P5 L51 need to provide a very brief description of a twilight board and how it is used, or else this statement is meaningless to the reader. P6 L3 The few seconds before fading are irrelevant if you tilt your head back and forth while looking at the sky and then use an object in the peripheral foreground (e.g. your arm) to point in the direction of the yellow brushes. P6 L11 You need to describe the 3 steps, otherwise the reader has no idea what you are referring to. Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 7 figures Figures 3 and 5 are confusing. I am well aware of the topic but cannot easily determine what these plots are showing Again I have no idea what Figure 7 is attempting to show, and the associated text in the manuscript P4 L41 does not help. Please write out in words what the statement you are attempting to make is and then use the figure to support that statement. This is important because you want your reader to understand your results and be able to use them, if not why bother publishing. Review form: Reviewer 2 (Barbara Webb) Is the manuscript scientifically sound in its present form? Yes Are the interpretations and conclusions justified by the results? Yes Is the language acceptable? Yes Is it clear how to access all supporting data? Yes Do you have any ethical concerns with this paper? No Have you any concerns about statistical analyses in this paper? No Recommendation? Accept with minor revision (please list in comments) Comments to the Author(s) This is an interesting paper that evaluates the hypothesis that Vikings could navigate using polarisation patterns visible to the naked eye. This is assessed by using a large database of recorded polarization levels in whole skies for different sun elevations and cloud cover percentage, and comparing these to previously established thresholds for perception of Haidinger's brushes. The title seems overlong, Perhaps’delectability of polarization cues for navigation under varied celestial conditions' would suffice? The final sentence of the abstract wcould be clearer it rearranged - ‘is most useful after sunset and prior to sunrise, when the sun is not visible and great sky' etc. line 55- l don't think the brushes would be well known to the typical reader. This is also why I suggest they are not featured in the title Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 8 p.2 line 10, it would be helpful to say a bit more about the thresholds. E-g. what percentile of tested subjects meet the most sensitive threshold? Or do they mean to say the lowest threshold measured for any subject was 23%, and the highest 87%? Is the distribution symmetric enough for the average to be meaningful? Briefly, how was the threshold determined? line 46, could mention the latitude of the measurements here (and somewhere in the discussion explain whether and how this might affect the polarisaton the final sentence of the introduction is uninformative, either remove or give an indication of the results. p3 liner 5, delete "from which we selected 1296 different skies". The grouping happens before the selection. It does not seem necessary to spell out all the ranges, or to use a numerical subscript rather than one that contains the relevant information. p4 line 3 (fig 3) I do not understand the description of this figure as plotting P(d*). It plots d against % sky with at least that value of d. The thresholds d* could be included in the figure, as vertical lines at the relevant valves. The same applies to fig 5. P4 line 28, if the elevation shown is 0 (as stated on line 16) then the elevation is not >0, so it is not clear what point is being made here. The sun is not visible in fig 4A. P5 lines 5 onward, it is not clear what information should be taken from the variance or delta_P. How might this affect the use of the method? P6 lines 7 onward, is it beyond the scope of this paper to carry out the error propagation steps to determine the practicality of the method? Decision letter (RSOS-160688) 04-Jan-2017 Dear Dr Horvath On behalf of the Editors, I am pleased to inform you that your Manuscript RSOS-160688 entitled "Sky-polarization prerequisite of Viking navigation with Haidinger’s brushes" has been accepted for publication in Royal Society Open Science subject to minor revision in accordance with the referee suggestions. Please find the referees' comments at the end of this email. The reviewers and handling editors have recommended publication, but also suggest some minor revisions to your manuscript. Therefore, I invite you to respond to the comments and revise your manuscript. 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Kind regards, Alice Power Editorial Coordinator Royal Society Open Science [email protected] on behalf of Jon Blundy Subject Editor, Royal Society Open Science [email protected] Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 11 Reviewer comments to Author: Reviewer: 1 Comments to the Author(s) Comments to Authors General Comments This is a great analysis of the proportion of the sky that would be useful for human navigation using celestial polarisation patterns relative to meteorological conditions. My only major concern with the manuscript is the comparison of the three different wavelength bands used. It is not necessary to provide all three colour bands, and in fact it detracts from the story by making it more complicated than it needs to be. The entoptics phenomenon of Haidinger's brushes is mediated by the absorption of short-wavelength light by the macular carotenoids (lutien, zeaxanthin and meso-zeaxanthin) (Müller et al 2016, and Bone 1984) and is not visible at wavelengths above about 500 nm, therefore the two colour bands in the green and red are not useful for perception of polarization and therefore do not add to your argument. Instead the extra data makes the story more confusing. If you remove the colour component you can focus on the main effect in the blue and make the story more concise and more easily accessible. In case you are interested, the original experiments we did (Temple et al. 2015) were done with humans looking at a modified LCD screen, however, I have since measured the human threshold using a sheet of rotating polarizer combined with a series of my custom made depolarizing filters and found that the human threshold is even lower than 23% we originally reported. Out of 20 people studied, four individuals could see HB down to 16% and one could go as low as 12%. In this study the mean d* was 32.4% and the min was 70%. If you want to add these results to this paper you could report them as unpublished data or personal communication. This work was done as part of a preliminary experiment that showed a correlation between threshold for detecting HB (d*) and macular pigment optical density (MPOD) measured on a dual wavelength scanning laser ophthalmoscope (Heidelberg Spectralis). These results concur with recent findings of Müller et al. 2016 (Müller PL, Müller S, Gliem M, et al. Perception of Haidinger brushes in macular disease depends on macular pigment density and visual acuity. Invest Ophthalmol Vis Sci. 2016;57:1448–1456. DOI:10.1167/ iovs.15-19004). It is not helpful for the reader when sentences have too many equations (e.g. P5 L5-11 ) or are written around a figure (e.g. start with "Figure x shows..."). It is much easier to read and understand the point being made if the sentences are written in words and just tell the story, leaving the equations and figures to the end to be put in parentheses e.g. Page 3 Line 56 "Figure 2 shows a totally overcast sky taken with our full-sky imaging polarimeter when the solar elevation was θ = 0º and the cloud percent was ρ = 100 %." It is much easier to understand if you focus on the results not the figure, as I have tried to do here "When skies are clear the proportion of the celestial hemisphere P suitable for human polarimetric navigation based on perception of Haidinger's brushes is high (insert P-value here; Figure 2). However, when the sky is overcast, P approaches 0 making it infeasible for navigation by this means (Fig. 3)." NOTE I would reverse the order in which these two points are presented and start with the positive situation in which polarimetric navigation IS possible rather than starting with when it IS NOT. Specific Comments To preclude, I really like this work and think that it will be a valuable contribution to the field but that it needs to be written in a way that makes it more accessible to the reader, so I have provided numerous suggestions below. Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 12 Page 1 Title The title is long and not easily understandable, specifically the expression "sky-polarization prerequisite" is not clear. Please consider an alternative, for example a variation of: Limits of human polarization perception support Viking navigation with the naked eye based on celestial polarization patterns. OR Celestial polarization patterns sufficient for Viking Navigation with Haidinger's brushes. OR Viking navigation with Haidinger's brushes: assessment of the meteorological conditions and proportion of the sky suitable based on human threshold for polarization perception. Page 1 Line 25 (P1 L25) Here 'brush' is used rather than "brushes" I recommend brushes but pick one and stick to it for consistency. P1 L25 you have already said '8-shaped' and I am not sure that 'brush-like' is a good description, I recall that the word brush was a poor translation of the German that was meant to be bushel, like a bushel of wheat. A better description is 'bowtie-like' rather than 'brush-like' P1 L25-28 This Sentence is long and difficult to understand with novel terminology that needs defining upon first use. "Since the celestial great circle passing through any sky point with high enough d-values and parallel to the yellow Haidinger’s brush crosses the sun, if two such sky points are selected, the sun position can be determined, which is at the intersection of the two great circles." specifically you need to define or come up with a better description of "the celestial great circle" And this statement is really unclear "passing through any sky point with high enough d-values and parallel to the yellow Haidinger’s brush crosses the sun" please see if you can reword this. The last part refers to "two great circles" but the first part makes it seem like there should only be one "the celestial great circle". Personally, I can see HB quite easily both in clear skies and in the forest when only a very small portion of the clear sky is visible through the canopy, and I don't think it is necessary to talk about 'great circles' but rather I think you could talk about a band of high degree of polarization that arcs across the sky at about 90 degrees to the position of the sun and that a person could pick two points on that band and observe the yellow brushes of HB and from that triangulate the position of the sun based on the orientation of those two observed brushes. But equally it does not need to be static observation. by sweeping your eye across the band of high DoP it is possible to get a dynamic view of HB that point directly at the sun. P1 L30 in addition to occlusion by cloud and fog, HB can also be used to locate the sun when it is below the horizon or occluded by objects on the horizon. P1 L30 the expression 'atmospheric optical prerequisite' is overly complicated, and this sentence could be simplified to something like "In order to determine the position of the sun using the celestial polarization pattern the d of the portion of the sky used, must be greater than the viewers degree of polarization threshold (d*) for perception of Haidinger's brushes." Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 13 P1 L34 remove the red and green here and throughout the paper (see general comment above) P1 L36 recommend changing 'cloud percent' to 'cloud cover (in percentage)' P1 L37 the statement 'We obtained that P is large enough only for the most sensitive human observers having perception threshold d*min = 23 % of Haidinger's brushes." does not make sense. "Large enough" for what? P1 L38 no space required between the number and the percent symbol, whereas there is a space required between a number and most of other units (e.g. nm) but not degree. P1 L39 recommend change to 'with cloud coverage of p = 0%, when the sun's position is already easily determined." delete "since it is visible with a big chance". P1 L42 again what does 'P is large enough' mean? if P is the proportion of the sky where d>d* then surely an observer only needs a couple of small areas of the sky that are larger the 5 degrees in radius (size of HB perception) to observe HB and detect the position of the sun, so if the d is high then only a small portion of the sky located near the band of maximum d is required. So what does 'large enough' mean? P1 L45 change 'great' to 'large' P1 L48 because on-line search engines will search your title and abstract it is redundant to use keywords that are already in your title and abstract in your list of keywords. Instead consider using other words that a potential reader might use to search for similar articles. e.g. calcite, orientation, sea-faring, orienteering, migration, vision, macula, macular pigment etc... P1 L53 see comment above about 'brush-like' could add that the two 8-shaped figures combine to form a Maltese cross shape P1 L54 & L55 stick to the '8-shaped figure' or perhaps 'bowtie-like shape' as used elsewhere P1 L55 delete 'well-known'. Haidinger's brushes are certainly not well known. I have been interested in this subject for near 20 years and very few vision scientists, optometrist, visual ecologist have ever heard of them and even few of the general population. P2 L2 change to 'brushes' here and throughout. P2 L2 suggest removing the word 'well' most people find it difficult to perceive HB at the best of times, and even those of us that can see them consistently would not say they are easy to see or are highly salient, it suffices to say that they can be perceived. P2 L4 suggest change to "when large portions of the sky are red and orange, or when the sky is..." P2 L11 correct 'treshold' to 'threshold' P2 L10 it is much easier to read if words are used rather than equations e.g. 'Recently, Temple et al [13] measured the degree of polarization threshold, d*, at which humans could just detect Haidinger's brushes. P2 L14 consider adding to this sentence as such "determined by the relative position of the observer, the sun and the celestial point observed (Fig. 1)." P2 L21 this 'celestial great circle' needs to be well defined earlier in the paper if it is going to be used. Otherwise consider 'band of highest polarization' Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 14 P2 L24 if is not clear why there are now 'two great circles'. I know what you are talking about but it is not written clearly enough for someone slightly unfamiliar with the specifics to understand it well enough. Furthermore there are no 'great circles' in your figure 1, nor should there be. I think it easier to describe it somewhat as I have in my comment above re P1 L25-28. P2 L34 change to 'they had their own analyser..." P2 L43 change to past tense 'the aim of this work was to study' P2 L44-45 remove red and green from here and the paper in general P2 L47 change to 'percent cloud cover' here and throughout the manuscript P2 L50 used past conditional past tense ' navigation could have been based on the perception...' P3 L14 change to 'coverage of the visible sky estimated by visual examination [40].' P3 L15 suggest 'refer to ever increasing cloud coverage' P3 L24 spelling 'approximately' P3 L30-32 this is a bit confusing, it is clear that there were 12 sun azimuth groups and 9 cloud coverage groups, but what are the '12 different skies"? Are these just 12 examples/samples of each of the above 108 groups? if so then this needs to me spelled out (described) more clearly. P3 L32-34 delete reference to colour P3 L43 consider 'probability that a viewer (Viking) with high, average and low sensitivity, respectively, ...' P3 L45 remove reference to colour P3 L50 the statement says that the areas that were not sky were masked out but they still show up in the figures (e.g. Fig 2 c,d, & e and 4 c,d & e) with red and blue regions suggesting under and overexposed regions, but really this just detracts from the message you are trying to get accross, so I recommend blacking out the non-sky regions in these images to remove the red a blue that is not in the sky (the only part of interest here). P3L57 see note in general comments about the order of presentation of results. Start with the positive situation i.e. clear skies (section 3.2 rather than 3.1) and use full sentences that do not start with "Figure X shows..." Actually by the time you remove the parts of 3.1 and 3.2 that belong in discussion and remove the reference to red and green the sections will be short enough that they can be combined into a single section that just focuses on percent cloud cover. P4 L9-12 this statement is interpretation of the results and therefore belongs in discussion section not results. P4 L16 change sentence so it does not start with "Figure 4A..." P4 L22-29 this statement is interpretation of the results and therefore belongs in discussion section not results. P4 L31 consider changing to "sun elevation-cloud cover matrix..." Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 15 P4 L33 change sentence so it does not start with "Figure 6..." P4 L44 change sentence so it says something more like "A person having low polarization sensitivity (e.g. d*max = 87%) can only detect HB under a select few meteorological conditions (Fig 6; Supplementary table S3, S6, S9) and would therefore be an unlikely choice for a Viking navigator." In the discussion, you could suggest that Viking navigators were likely those that had the highest polarization sensitivity (So move P4 L59 to discussion), you could also add that a diet rich in fish, which is a good source of macular carotenoids, may have contributed to Viking's general ability to see HB. P4 L50 same as above. P5L5-11 this paragraph is almost incomprehensible, it is written as an equation masquerading as a sentence. Rewrite as a sentence with words rather than symbols. P5 L20 This paragraph starts off well with clear words used to describe everything and then the reader gets to this line that starts with <P>(θ,ρ), and gets totally lost. Please write out in words what these symbols mean as you did in the first two sentences. P5 L25 need to change this to blue since polarization sensitivity has nothing to do with the human spectral sensitivity curve that peaks in green, and everything to do with the polarization sensitivity, which is maximal at 460 nm (2nd sentence of Chapter 14 of your book Gábor that we wrote together) due to absorption of macular pigments (Bone 1980 Vision research 20:213-220). The statement that blue can be ignored is incorrect. In blue light the HB become clearer, and it is thought that the yellow colour a person sees under full spectrum illumination is a psychophysical effect of the eye where yellow is perceived when there is an absence of blue. I agree that the blue brushes of HB are or little utility for navigation but that has nothing to do with the spectral content of the sky. P5 L31 & 34 again consider rewording the 'great circle passing the sun', especially the last part 'passing the sun' this does not make sense. P5 L35-45 suggest changing to something like... "We found that the proportion of the sky suitable for the humans with a high sensitivity to Haidinger's brushes (d*min = 23%) is high under numerous meteorological conditions, particularly when the sun is low on the horizon and when the sky is clear of cloud. When the sun is below the horizon, and therefore not possible to see or accurately locate, the proportion of the sky suitable for sky polarimetric navigation is between 40% and 77% depending on cloud cover. P5 L48 Need to define "3rd step" of just use the word-based description first and then put '3rd step' in brackets and state that this is what they called it. P5 L51 need to provide a very brief description of a twilight board and how it is used, or else this statement is meaningless to the reader. P6 L3 The few seconds before fading are irrelevant if you tilt your head back and forth while looking at the sky and then use an object in the peripheral foreground (e.g. your arm) to point in the direction of the yellow brushes. P6 L11 You need to describe the 3 steps, otherwise the reader has no idea what you are referring to. figures Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 16 Figures 3 and 5 are confusing. I am well aware of the topic but cannot easily determine what these plots are showing Again I have no idea what Figure 7 is attempting to show, and the associated text in the manuscript P4 L41 does not help. Please write out in words what the statement you are attempting to make is and then use the figure to support that statement. This is important because you want your reader to understand your results and be able to use them, if not why bother publishing. Reviewer: 2 Comments to the Author(s) This is an interesting paper that evaluates the hypothesis that Vikings could navigate using polarisation patterns visible to the naked eye. This is assessed by using a large database of recorded polarization levels in whole skies for different sun elevations and cloud cover percentage, and comparing these to previously established thresholds for perception of Haidinger's brushes. The title seems overlong, Perhaps’delectability of polarization cues for navigation under varied celestial conditions' would suffice? The final sentence of the abstract wcould be clearer it rearranged - ‘is most useful after sunset and prior to sunrise, when the sun is not visible and great sky' etc. line 55- l don't think the brushes would be well known to the typical reader. This is also why I suggest they are not featured in the title p.2 line 10, it would be helpful to say a bit more about the thresholds. E-g. what percentile of tested subjects meet the most sensitive threshold? Or do they mean to say the lowest threshold measured for any subject was 23%, and the highest 87%? Is the distribution symmetric enough for the average to be meaningful? Briefly, how was the threshold determined? line 46, could mention the latitude of the measurements here (and somewhere in the discussion explain whether and how this might affect the polarisaton the final sentence of the introduction is uninformative, either remove or give an indication of the results. p3 liner 5, delete "from which we selected 1296 different skies". The grouping happens before the selection. It does not seem necessary to spell out all the ranges, or to use a numerical subscript rather than one that contains the relevant information. p4 line 3 (fig 3) I do not understand the description of this figure as plotting P(d*). It plots d against % sky with at least that value of d. The thresholds d* could be included in the figure, as vertical lines at the relevant valves. The same applies to fig 5. Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 17 P4 line 28, if the elevation shown is 0 (as stated on line 16) then the elevation is not >0, so it is not clear what point is being made here. The sun is not visible in fig 4A. P5 lines 5 onward, it is not clear what information should be taken from the variance or delta_P. How might this affect the use of the method? P6 lines 7 onward, is it beyond the scope of this paper to carry out the error propagation steps to determine the practicality of the method? Author's Response to Decision Letter for (RSOS-160688) See Appendix A. Decision letter (RSOS-160688.R1) 09-Jan-2017 Dear Dr Horvath, I am pleased to inform you that your manuscript entitled "Sky-polarimetric Viking navigation with Haidinger’s brushes" is now accepted for publication in Royal Society Open Science. You can expect to receive a proof of your article in the near future. Please contact the editorial office ([email protected] and [email protected]) to let us know if you are likely to be away from e-mail contact. Due to rapid publication and an extremely tight schedule, if comments are not received, your paper may experience a delay in publication. Royal Society Open Science operates under a continuous publication model (http://bit.ly/cpFAQ). Your article will be published straight into the next open issue and this will be the final version of the paper. As such, it can be cited immediately by other researchers. As the issue version of your paper will be the only version to be published I would advise you to check your proofs thoroughly as changes cannot be made once the paper is published. In order to raise the profile of your paper once it is published, we can send through a PDF of your paper to selected colleagues. If you wish to take advantage of this, please reply to this email with the name and email addresses of up to 10 people who you feel would wish to read your article. On behalf of the Editors of Royal Society Open Science, we look forward to your continued contributions to the Journal. Best wishes, Alice Power Editorial Coordinator Royal Society Open Science [email protected] Appendix A Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Dear Editorial Office of the Royal Soceity Open Science, I would like to submit our revised manuscript RSOS-160688_R1 entitled Celestial polarization patterns sufficient for Viking navigation with the naked eye: detectability of Haidinger's brushes on the sky versus meteorological conditions by Gabor Horvath, Peter Takacs, Balazs Kretzer, Szilvia Szilasi, Denes Szaz, Alexandra Farkas, Andras Barta to the Royal Society Open Science as a research article. Our manuscript was revised on the basis of the reports of two Reviewers. All relevant changes performed on the basis of the comments of Referee 1 and Referee 2 are marked by violet and green, respectively. Here we give our response to the points of Reviewer 1. Our Point-by-Point Response to the Comments of Reviewers 1 and 2 can be found below. Sincerely yours: Gabor Horvath (corresponding author) Dr. Gabor Horvath (D.Sc.) Environmental Optics Laboratory, Department of Biological Physics, Physical Institute, Eötvös University, H-1117 Budapest, Pazmany setany 1, Hungary e-mail: [email protected] http://arago.elte.hu Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Point-by-Point Response to the Comments of Reviewer 1 We thank for the positive and constructive comments of Referee 1. Our manuscript was revised on the basis of the reports of two Reviewers. All relevant changes performed on the basis of the comments of Referee 1 and Referee 2 are marked by violet and green, respectively. Here we give our response to the points of Reviewer 1. Referee 1 wrote: My only major concern with the manuscript is the comparison of the three different wavelength bands used. It is not necessary to provide all three colour bands, and in fact it detracts from the story by making it more complicated than it needs to be. The entoptics phenomenon of Haidinger's brushes is mediated by the absorption of short-wavelength light by the macular carotenoids (lutien, zeaxanthin and meso-zeaxanthin) (Müller et al 2016, and Bone 1984) and is not visible at wavelengths above about 500 nm, therefore the two colour bands in the green and red are not useful for perception of polarization and therefore do not add to your argument. Instead the extra data makes the story more confusing. If you remove the colour component you can focus on the main effect in the blue and make the story more concise and more easily accessible. Answer: As suggested, in our revised paper we limited our study onto the blue (450 nm) spectral range. The text, tables and figures were changed accordingly. Referee 1 wrote: In case you are interested, the original experiments we did (Temple et al. 2015) were done with humans looking at a modified LCD screen, however, I have since measured the human threshold using a sheet of rotating polarizer combined with a series of my custom made depolarizing filters and found that the human threshold is even lower than 23% we originally reported. Out of 20 people studied, four individuals could see HB down to 16% and one could go as low as 12%. In this study the mean d* was 32.4% and the min was 70%. If you want to add these results to this paper you could report them as unpublished data or personal communication. This work was done as part of a preliminary experiment that showed a correlation between threshold for detecting HB (d*) and macular pigment optical density (MPOD) measured on a dual wavelength scanning laser ophthalmoscope (Heidelberg Spectralis). These results concur with recent findings of Müller et al. 2016 (Müller PL, Müller S, Gliem M, et al. Perception of Haidinger brushes in macular disease depends on macular pigment density and visual acuity. Invest Ophthalmol Vis Sci. 2016;57:1448– 1456. DOI:10.1167/ iovs.15-19004). Answer: Your preliminary results mentioned above were not included into our revised manuscript. We wait for their publication. Referee 1 wrote: It is not helpful for the reader when sentences have too many equations (e.g. P5 L511 ) or are written around a figure (e.g. start with "Figure x shows..."). It is much easier to read and understand the point being made if the sentences are written in words and just tell the story, leaving the equations and figures to the end to be put in parentheses e.g. Page 3 Line 56 "Figure 2 shows a totally overcast sky taken with our full-sky imaging polarimeter when the solar elevation was θ = 0º and the cloud percent was ρ = 100 %." It is much easier to understand if you focus on the results not the figure, as I have tried to do here "When skies are clear the proportion of the celestial hemisphere P suitable for human polarimetric navigation based on perception of Haidinger's brushes is high (insert P-value here; Figure 2). However, when the sky is overcast, P approaches 0 making it infeasible for navigation by this means (Fig. 3)." NOTE I would reverse the order in which these two points are presented and start with the positive situation in which polarimetric navigation IS possible rather than starting with when it IS NOT. Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Answer: The order of overcast and clear skies was reversed, and the text was changed as suggested. Referee 1 wrote: Page 1 Title: The title is long and not easily understandable, specifically the expression "sky-polarization prerequisite" is not clear. Please consider an alternative, for example a variation of: Limits of human polarization perception support Viking navigation with the naked eye based on celestial polarization patterns OR Celestial polarization patterns sufficient for Viking Navigation with Haidinger's brushes OR Viking navigation with Haidinger's brushes: assessment of the meteorological conditions and proportion of the sky suitable based on human threshold for polarization perception Answer: Taking into consideration the suggestions of both Referees, the new title is: Celestial polarization patterns sufficient for Viking navigation with the naked eye: detectability of Haidinger's brushes on the sky versus meteorological conditions Referee 1 wrote: Page 1 Line 25 (P1 L25) Here 'brush' is used rather than "brushes". I recommend brushes but pick one and stick to it for consistency. P1 L25 you have already said '8-shaped' and I am not sure that 'brush-like' is a good description, I recall that the word brush was a poor translation of the German that was meant to be bushel, like a bushel of wheat. A better description is 'bowtie-like' rather than 'brush-like' Answer: These changes were done in the revised manuscript. Referee 1 wrote: P1 L25-28 This Sentence is long and difficult to understand with novel terminology that needs defining upon first use. "Since the celestial great circle passing through any sky point with high enough d-values and parallel to the yellow Haidinger’s brush crosses the sun, if two such sky points are selected, the sun position can be determined, which is at the intersection of the two great circles." Specifically you need to define or come up with a better description of "the celestial great circle". And this statement is really unclear "passing through any sky point with high enough d-values and parallel to the yellow Haidinger’s brush crosses the sun" please see if you can reword this. The last part refers to "two great circles" but the first part makes it seem like there should only be one "the celestial great circle". Personally, I can see HB quite easily both in clear skies and in the forest when only a very small portion of the clear sky is visible through the canopy, and I don't think it is necessary to talk about 'great circles' but rather I think you could talk about a band of high degree of polarization that arcs across the sky at about 90 degrees to the position of the sun and that a person could pick two points on that band and observe the yellow brushes of HB and from that triangulate the position of the sun based on the orientation of those two observed brushes. But equally it does not need to be static observation. by sweeping your eye across the band of high DoP it is possible to get a dynamic view of HB that point directly at the sun. Answer: This sentence was rewritten as follows: ’A band of high d arcs across the sky at 90o from the sun. A person can pick two points on that band, observe the yellow brushes, and triangulate the position of the sun based on the orientation of the two observed brushes.’ Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Referee 1 wrote: P1 L30 in addition to occlusion by cloud and fog, HB can also be used to locate the sun when it is below the horizon or occluded by objects on the horizon. Answer: The following sentence was here added: ’Furthermore, Haidinger’s brushes can also be used to locate the sun when it is below the horizon or occluded by objects on the horizon.’ Referee 1 wrote: P1 L30 the expression 'atmospheric optical prerequisite' is overly complicated, and this sentence could be simplified to something like "In order to determine the position of the sun using the celestial polarization pattern the d of the portion of the sky used, must be greater than the viewers degree of polarization threshold (d*) for perception of Haidinger's brushes." Answer: This change was performed. Referee 1 wrote: P1 L34 remove the red and green here and throughout the paper (see general comment above) P1 L36 recommend changing 'cloud percent' to 'cloud cover (in percentage)' Answer: As suggested, red and green data were deleted throughout the paper. Referee 1 wrote: P1 L37 the statement 'We obtained that P is large enough only for the most sensitive human observers having perception threshold d*min = 23 % of Haidinger's brushes." does not make sense. "Large enough" for what? Answer: This sentence was deleted. Referee 1 wrote: P1 L38 no space required between the number and the percent symbol, whereas there is a space required between a number and most of other units (e.g. nm) but not degree. Answer: These changes were done throughout the revised manuscript.. Referee 1 wrote: P1 L39 recommend change to 'with cloud coverage of p = 0%, when the sun's position is already easily determined." delete "since it is visible with a big chance". Answer: These changes were done. Referee 1 wrote: P1 L42 again what does 'P is large enough' mean? If P is the proportion of the sky where d>d* then surely an observer only needs a couple of small areas of the sky that are larger the 5 degrees in radius (size of HB perception) to observe HB and detect the position of the sun, so if the d is high then only a small portion of the sky located near the band of maximum d is required. So what does 'large enough' mean? Answer: This sentence was rewritten as follows: ’If the sun is below the horizon (–5o ≤ θ < 0°) during twilight, P = 76.17 ± 4.18% for d*min = 23% under clear sky conditions.’ Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Referee 1 wrote: P1 L45 change 'great' to 'large' Answer: This change was done. Referee 1 wrote: P1 L48 because on-line search engines will search your title and abstract it is redundant to use keywords that are already in your title and abstract in your list of keywords. Instead consider using other words that a potential reader might use to search for similar articles. e.g. calcite, orientation, sea-faring, orienteering, migration, vision, macula, macular pigment etc... Answer: We kept the original keywords, because the suggested ones are not typical for the content of our paper. However, we added the keyword human polarization vision. Referee 1 wrote: P1 L53 see comment above about 'brush-like'. Could add that the two 8-shaped figures combine to form a Maltese cross shape P1 L54 & L55 stick to the '8-shaped figure' or perhaps 'bowtie-like shape' as used elsewhere Answer: These changes were done. Referee 1 wrote: P1 L55 delete 'well-known'. Haidinger's brushes are certainly not well known. I have been interested in this subject for near 20 years and very few vision scientists, optometrist, visual ecologist have ever heard of them and even few of the general population. Answer: In this sentence ’well-known’ was replaced by ’so-called’. Referee 1 wrote: P2 L2 change to 'brushes' here and throughout. P2 L2 suggest removing the word 'well' most people find it difficult to perceive HB at the best of times, and even those of us that can see them consistently would not say they are easy to see or are highly salient, it suffices to say that they can be perceived. P2 L4 suggest change to "when large portions of the sky are red and orange, or when the sky is..." P2 L11 correct 'treshold' to 'threshold' P2 L10 it is much easier to read if words are used rather than equations e.g. 'Recently, Temple et al [13] measured the degree of polarization threshold, d*, at which humans could just detect Haidinger's brushes. P2 L14 consider adding to this sentence as such "determined by the relative position of the observer, the sun and the celestial point observed (Fig. 1)." Answer: All these changes were done. Referee 1 wrote: P2 L21 this 'celestial great circle' needs to be well defined earlier in the paper if it is going to be used. Otherwise consider 'band of highest polarization' P2 L24 if is not clear why there are now 'two great circles'. I know what you are talking about but it is not written clearly enough for someone slightly unfamiliar with the specifics to understand it well Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 enough. Furthermore there are no 'great circles' in your figure 1, nor should there be. I think it easier to describe it somewhat as I have in my comment above re P1 L25-28. Answer: Referee 1 mistakes here two quite different concepts: The band of highest degrees of polarization is at 90o from the sun, and has nothing to do with a spherical geometric celestial great circle passing through a given sky point. The concept ’great circle’ is used only in this paragraph, and its meaning should be clear in Figure 1. Referee 1 wrote: P2 L34 change to 'they had their own analyser..." P2 L43 change to past tense 'the aim of this work was to study' P2 L44-45 remove red and green from here and the paper in general P2 L47 change to 'percent cloud cover' here and throughout the manuscript P3 L14 change to 'coverage of the visible sky estimated by visual examination [40].' P3 L15 suggest 'refer to ever increasing cloud coverage' P3 L24 spelling 'approximately' Answer: All these changes were done. Referee 1 wrote: P3 L30-32 this is a bit confusing, it is clear that there were 12 sun azimuth groups and 9 cloud coverage groups, but what are the '12 different skies"? Are these just 12 examples/samples of each of the above 108 groups? If so then this needs to me spelled out (described) more clearly. Answer: This sentence was changed as follows: ’Thus, we created 12 × 9 = 108 (θ, ρ) groups. In each group we selected 12 different sky samples (with θ- and ρ-values falling into the θ- and ρ-interval of the given group) from our polarimetric sky archives.’ Referee 1 wrote: P3 L32-34 delete reference to colour P3 L43 consider 'probability that a viewer (Viking) with high, average and low sensitivity, respectively, ...' P3 L45 remove reference to colour Answer: Done. Referee 1 wrote: P3 L50 the statement says that the areas that were not sky were masked out but they still show up in the figures (e.g. Fig 2 c,d, & e and 4 c,d & e) with red and blue regions suggesting under and overexposed regions, but really this just detracts from the message you are trying to get accross, so I recommend blacking out the non-sky regions in these images to remove the red a blue that is not in the sky (the only part of interest here). Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Answer: In the patterns of the degree of polarization d, black means d = 100%, thus the non-sky regions as well as the under- and overexposed celestial areas cannot be blacked out, otherwise the reader could think that these regions and areas are also maximally polarized. Thus, we kept the original d-patterns with blue (underexposed) and red (overexposed) colours. This is our traditional colour code used in our numerous publications presenting polarization patterns. Referee 1 wrote: P3 L57 see note in general comments about the order of presentation of results. Start with the positive situation i.e. clear skies (section 3.2 rather than 3.1) and use full sentences that do not start with "Figure X shows..." Actually by the time you remove the parts of 3.1 and 3.2 that belong in discussion and remove the reference to red and green the sections will be short enough that they can be combined into a single section that just focuses on percent cloud cover. P4 L9-12 this statement is interpretation of the results and therefore belongs in discussion section not results. P4 L16 change sentence so it does not start with "Figure 4A..." P4 L22-29 this statement is interpretation of the results and therefore belongs in discussion section not results. P4 L31 consider changing to "sun elevation-cloud cover matrix..." P4 L33 change sentence so it does not start with "Figure 6..." P4 L44 change sentence so it says something more like "A person having low polarization sensitivity (e.g. d*max = 87%) can only detect HB under a select few meteorological conditions (Fig 6; Supplementary table S3, S6, S9) and would therefore be an unlikely choice for a Viking navigator." In the discussion, you could suggest that Viking navigators were likely those that had the highest polarization sensitivity (So move P4 L59 to discussion), you could also add that a diet rich in fish, which is a good source of macular carotenoids, may have contributed to Viking's general ability to see HB. P4 L50 same as above. P5 L5-11 this paragraph is almost incomprehensible, it is written as an equation masquerading as a sentence. Rewrite as a sentence with words rather than symbols. P5 L20 This paragraph starts off well with clear words used to describe everything and then the reader gets to this line that starts with <P>(θ,ρ), and gets totally lost. Please write out in words what these symbols mean as you did in the first two sentences. Answer: These changes were done. Referee 1 wrote: P5 L25 need to change this to blue since polarization sensitivity has nothing to do with the human spectral sensitivity curve that peaks in green, and everything to do with the polarization sensitivity, which is maximal at 460 nm (2nd sentence of Chapter 14 of your book Gábor that we wrote together) due to absorption of macular pigments (Bone 1980 Vision research 20:213- Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 220). The statement that blue can be ignored is incorrect. In blue light the HB become clearer, and it is thought that the yellow colour a person sees under full spectrum illumination is a psychophysical effect of the eye where yellow is perceived when there is an absence of blue. I agree that the blue brushes of HB are or little utility for navigation but that has nothing to do with the spectral content of the sky. P5 L31 & 34 again consider rewording the 'great circle passing the sun', especially the last part 'passing the sun' this does not make sense. Answer: This paragraph was revised in the following way: ’In the analysis of the proportion P of the sky for which the degree d of sky polarization is larger than the perception threshold d* (= 23, 56, 87%), the most important spectral range is the blue, in which the human eye perceives Haidinger’s brushes with a maximal sensitivity at 460 nm due to absorption of macular pigments [39]. In blue light the Haidinger’s brushes become clearer, and it is thought that the yellow colour a person sees under full spectrum illumination is a psychophysical effect of the eye where yellow is perceived when there is an absence of blue [1,2]. Thus, our study was restricted to this part of the spectrum. We obtained that the mean <P> of P is large only for the most sensitive-eyed human observers (Viking navigators) having a perception threshold d*min = 23% of Haidinger’s brushes. Viking navigators were likely those that had the highest polarization sensitivity (i.e. having the lowest threshold d*). A diet rich in fish, which is a good source of macular carotenoids, may have contributed to Viking's general ability to see Haidinger’s brushes (Shelby Temple, personal communication).’ Referee 1 wrote: P5 L35-45 suggest changing to something like... "We found that the proportion of the sky suitable for the humans with a high sensitivity to Haidinger's brushes (d*min = 23%) is high under numerous meteorological conditions, particularly when the sun is low on the horizon and when the sky is clear of cloud. When the sun is below the horizon, and therefore not possible to see or accurately locate, the proportion of the sky suitable for sky polarimetric navigation is between 40% and 77% depending on cloud cover. Answer: This change was done. Referee 1 wrote: P5 L48 Need to define "3rd step" of just use the word-based description first and then put '3rd step' in brackets and state that this is what they called it. Answer: This sentence was revised as follows: ’This finding corresponds to the results of Száz et al. [38], who also found that regarding the estimation of solar elevation angle with numbers of fist and finger of outstretched arms in the the third step of skypolarimetric Viking navigation, the most appropriate time for navigation is around sunset and sunrise.’ Referee 1 wrote: P5 L51 need to provide a very brief description of a twilight board and how it is used, or else this statement is meaningless to the reader. Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Answer: All details of the rather complicated use of a twilight board, a shadow-stick and sunstone crystals are described in the cited paper of Bernáth et al. [31]. The only important issue here is that this method functions well at twilight. Thus, we omitted a very brief description of a twilight board and how it is used. [31] Bernáth B, Farkas A, Száz D, Blahó M, Egri Á, Barta A, Åkesson S, Horváth G (2014) How could the Viking sun compass be used with sunstones before and after sunset? Twilight board as a new interpretation of the Uunartoq artefact fragment. Proceedings of the Royal Society A 470: article number 20130787 This paragraph was slightly rewritten as follows: ’The navigation method deduced by Bernáth et al. [31] based on the Uunartoq artefact fragment functioning as a twilight board is useful also during twilight and under clear skies, when the sun is close to the horizon (when −8° ≤ θ ≤ +10°). Using a twilight board, a shadow-stick and sunstone crystals could have allowed Vikings to navigate during long twilight periods on the basis of polarization patterns of clear skies. Based on our results in this paper, we can conclude, that birefringent (e.g. calcite) or dichroic (e.g. tourmaline or cordierite) sunstone crystals could be replaced with Haidinger’s brushes in the sky-polarimetric Viking navigation.’ Referee 1 wrote: P6 L3 The few seconds before fading are irrelevant if you tilt your head back and forth while looking at the sky and then use an object in the peripheral foreground (e.g. your arm) to point in the direction of the yellow brushes. Answer: The criticized sentences were revised in the following way: ’For example, it is questionable whether the few seconds prior to fading of the Haidinger’s brushes when looking at the polarized sky with a fixed head are enough to determine the position of the occluded sun in the sky. However, this few seconds before fading are irrelevant, if our head is tilted periodically back and forth while looking at the sky and then we use an object in the peripheral foreground (e.g. our arm) to point in the direction of the yellow Haidinger’s brushes. Nevertheless, it would be worth studying the accuracy and reliability of Viking navigation based on Haidinger’s brushes under real open air conditions.’ Referee 1 wrote: P6 L11 You need to describe the 3 steps, otherwise the reader has no idea what you are referring to. Answer: This sentence was rewritten as follows: ’Such computations have been presented by Farkas et al. [34] and Száz et al. [37,38] for this navigation method using dichroic and birefringent sunstone crystals.’ Referee 1 wrote: Figures 3 and 5 are confusing. I am well aware of the topic but cannot easily determine what these plots are showing Answer: The original Figs. 3 and 5 were merged to the present Fig. 3 with a more understandable legend. Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Referee 1 wrote: Again I have no idea what Figure 7 is attempting to show, and the associated text in the manuscript P4 L41 does not help. Please write out in words what the statement you are attempting to make is and then use the figure to support that statement. This is important because you want your reader to understand your results and be able to use them, if not why bother publishing. Answer: About Fig. 4B (originally Fig. 7) in our revised manuscript we wrote: ’The most sensitive persons (d*min = 23%) possess the widest range of the standard deviation (1.97% ≤ ΔP ≤ 29.66%) for medium percents of cloud cover ρ ≤ 75% and solar elevations not larger than 35º (Fig. 4B, Table 4). Persons with an average sensitivity (d*ave = 56%) have small standard deviations (3.14% ≤ ΔP ≤ 10.26%) for smaller percents of cloud cover ρ ≤ 25% and lower solar elevations θ ≤ 15º (Fig. 4B, Table 5). For the least polarization-sensitive persons (with d*max = 87%) the standard deviations of <P> are practically zero (ΔP ≤ 0.2681%; Fig. 4B, Table 6). The larger the standard deviation ΔP at a given meteorological situation (θ-ρ cell), the smaller the reliability of the navigation method based on Haidinger’s brushes under that sky conditions.’ Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Point-by-Point Response to the Comments of Reviewer 2 We thank for the positive and constructive comments of Referee 2. Our manuscript was revised on the basis of the reports of two Reviewers. All relevant changes performed on the basis of the comments of Referee 1 and Referee 2 are marked by violet and green, respectively. Here we give our response to the points of Reviewer 2. Referee 2 wrote: The title seems overlong, Perhaps ’detectability of polarization cues for navigation under varied celestial conditions' would suffice? Answer: Taking into consideration the suggestions of both Referees, the new title is: Celestial polarization patterns sufficient for Viking navigation with the naked eye: detectability of Haidinger's brushes on the sky versus meteorological conditions Referee 2 wrote: The final sentence of the abstract could be clearer it rearranged - ‘is most useful after sunset and prior to sunrise, when the sun is not visible and great sky' etc. Answer: This sentence was rewritten as follows: ’Consequently, the sky-polarimetric Viking navigation based on Haidinger’s brushes is most useful after sunset and prior to sunrise, when the sun is not visible and large sky regions are bright, clear and polarized enough for perception of Haidinger’s brushes.’ Referee 2 wrote: line 55, I don't think the brushes would be well known to the typical reader. This is also why I suggest they are not featured in the title. Answer: The criticized word ’well-known’ was replaced by so-called: ’These are the so-called Haidinger’s brushes, the angular extension of which is about 5o centered at the fovea of the human retina, and are mediated by dichroic carotenoids in the macula lutea [1-7].’ However, we kept ’Haidinger’s brushes’ in the title, because they are the main subject of this paper. Referee 2 wrote: p.2 line 10, it would be helpful to say a bit more about the thresholds. E-g. what percentile of tested subjects meet the most sensitive threshold? Or do they mean to say the lowest threshold measured for any subject was 23%, and the highest 87%? Is the distribution symmetric enough for the average to be meaningful? Briefly, how was the threshold determined? Answer: As requested, in the revised Introduction we added the following: ’Recently, Temple et al. [13] measured the degree of polarization threshold d*, at which humans could just detect Haidinger's brushes. They presented gratings in polarization-only contrast at varying degrees of polarization d in order to measure the lower limits of human polarized light detection. Participants were, on average, able to perform the task down to a threshold of d*ave = 56 ± 3%, and there was a normal continuous distribution of threshold values extending down to as low as d*min = 23% and up to as high as d*max = 87% [13].’ Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Referee 2 wrote: line 46, could mention the latitude of the measurements here and somewhere in the discussion explain whether and how this might affect the polarisaton. Answer: As requested, in the revised Introduction we wrote: ’Using full-sky imaging polarimetry, we measured the d-pattern of skylight in the blue (450 nm) spectral range for 1296 different meteorological conditions with different solar elevation angles θ and percent cloud cover ρ in Hungary (at latitude 47° 15' 29.83” N).’ In the revised Discussion we wrote: ’Although we measured the sky polarization for 1296 meteorological situations with different solar elevations θ and cloudinesses ρ in Hungary at latitude 47° 15' 29.83” N, this might not affect considerably sky polarizaton, because for a given ρ, celestial polarization depends predominantly on θ, and we restricted our measurements to solar elevations –5o ≤ θ ≤ +55o occurring at 61o latitude where the main Viking sailing route ran. The range 0 okta ≤ ρ ≤ 8 oktas used by us covered the whole range of cloudiness that can also occur at 61o latitude.’ Referee 2 wrote: The final sentence of the introduction is uninformative, either remove or give an indication of the results. Answer: As suggested, this sentence was deleted. Referee 2 wrote: p3 line 5, delete "from which we selected 1296 different skies". The grouping happens before the selection. It does not seem necessary to spell out all the ranges, or to use a numerical subscript rather than one that contains the relevant information. Answer: The criticized sentence part was deleted. Referee 2 wrote: p4 line 3 (fig 3) I do not understand the description of this figure as plotting P(d*). It plots d against % sky with at least that value of d. The thresholds d* could be included in the figure, as vertical lines at the relevant values. The same applies to fig 5. Answer: The legend of Figure 3 (being merged with Fig. 5) was revised as suggested: ’Figure 3: Proportion P (%) of the clear and overcast skies in Fig. 2 with degrees of skylight polarization not lower than d in the blue (450 nm) spectral range. The vertical lines show the values d* = 23, 56, 87% being the three characteristic perception thresholds of Haidinger’s brushes.’ Referee 2 wrote: P4 line 28, if the elevation shown is 0 (as stated on line 16) then the elevation is not > 0, so it is not clear what point is being made here. The sun is not visible in fig 4A. Answer: Referee 2 is right, therefore the criticized following sentence was deleted: ’However, in this case this would have been unnecessary, because the sun is visible under clear sky conditions, if the solar elevation θ > 0º.’ Referee 2 wrote: P5 lines 5 onward, it is not clear what information should be taken from the variance or delta_P. How might this affect the use of the method? Downloaded from http://rsos.royalsocietypublishing.org/ on June 17, 2017 Answer: At the end the Results section we added: ’The larger the standard deviation ΔP at a given meteorological situation (θ-ρ cell), the smaller the reliability of the navigation method based on Haidinger’s brushes under that sky conditions.’ Referee 2 wrote: P6 lines 7 onward, is it beyond the scope of this paper to carry out the error propagation steps to determine the practicality of the method? Answer: At the end of the revised Discussion we wrote: ’It is beyond the scope of this paper to carry out such error propagation to determine the practicality of the navigation method based on Haidinger’s brushes. This can be an interesting task of future research.’
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