The DOI matches its Crossref record; 22 of the 22 references it cites were found in a registry or an index.
Experimental determination of refractive index of condensed reflectin in squid iridocytes
Open accessAbstract
Loliginid squid dynamically tune the structural iridescence of cells in their skin for active camouflage and communication. Bragg reflectors in these cells consist of membrane-bound lamellae periodically alternating with low refractive index extracellular spaces; neuronal signalling induces condensation of the reflectin proteins that fill the lamellae, consequently triggering the expulsion of water. This causes an increase in refractive index within the lamellae, activating reflectance, with the change in lamellar thickness and spacing progressively shifting the wavelength of reflected light. We used micro-spectrophotometry to measure the functionally relevant refractive index of the high-index lamellae of the Bragg reflectors containing the condensed reflectins in chemically fixed dermal iridocytes of the squid,Doryteuthis opalescens. Our high-magnification imaging spectrometer allowed us to obtain normalized spectra of optically distinct sections of the individual, subcellular, multi-layer Bragg stacks. Replacement of the extracellular fluid with liquids of increasing refractive index allowed us to measure the reflectivity of the Bragg stacks as it decreased progressively to 0 when the refractive index of the extracellular medium exactly matched that of the reflectin-filled lamellae, thus allowing us to directly measure the refractive index of the reflectin-filled lamellae as ncondensed lamellae≈ 1.44. The measured value of the physiologically relevant ncondensed lamellae from these bright iridocytes falls within the range of values that we recently determined by an independent optical method and is significantly lower than values previously reported for dehydrated and air-dried reflectin films. We propose that this directly measured value for the refractive index of the squid's Bragg lamellae containing the condensed reflectins is most appropriate for calculations of reflectivity in similar reflectin-based high-index layers in other molluscs.
Overview
Integrity signals
Registry
Cites
Statistics
Verification
Claims the paper makes
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Structural coloration is observed without clear underlying ultrastructural electron-dense reflectin-filled lamellae similar to the signature plate-like stacks of cephalopod iridocytes.
“One of the most interesting aspects of this system is that structural coloration is observed without clear underlying ultrastructural electron-dense reflectin-filled lamellae similar to the signature plate-like stacks of cephalopod iridocytes7,41.”
From Dynamic pigmentary and structural coloration within cephalopod chromatophore organs (2019) · cites this paper as [41]
Citations
Cites
The physics and biology of animal reflectors
Land, M. F. (1972) · Progress in Biophysics and Molecular Biology · doi:10.1016/0079-6107(72)90004-1 · Cited by 598
Mechanisms and behavioural functions of structural coloration in cephalopods
Mäthger, L. M., Denton, E. J., Marshall, N. J., & Hanlon, R. T. (2008) · Journal of The Royal Society Interface · doi:10.1098/rsif.2008.0366.focus · Cited by 347
Reflectins: The Unusual Proteins of Squid Reflective Tissues
Crookes, W. J., Ding, L., Huang, Q., Kimbell, J. R., Horwitz, J., & McFall‐Ngai, M. (2004) · Science · doi:10.1126/science.1091288 · Cited by 228
Mechanism of reflexion in silvery layers of fish and cephalopods
Denton, E. J., & Land, M. F. (1971) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.1971.0051 · Cited by 227
The self-organizing properties of squid reflectin protein
Kramer, R. M., Crookes-Goodson, W. J., & Naik, R. R. (2007) · Nature Materials · doi:10.1038/nmat1930 · Cited by 172
Mäthger, L. M., & Hanlon, R. T. (2007) · Cell and Tissue Research · doi:10.1007/s00441-007-0384-8 · Cited by 171
Chromatophore Organs, Reflector Cells, Iridocytes and Leucophores in Cephalopods
CLONEY, R. A., & BROCCO, S. L. (1983) · American Zoologist · doi:10.1093/icb/23.3.581 · Cited by 129
DeMartini, D. G., Krogstad, D. V., & Morse, D. E. (2013) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1217260110 · Cited by 124
Self-citationBright White Scattering from Protein Spheres in Color Changing, Flexible Cuttlefish Skin
Mäthger, L. M., Senft, S. L., Gao, M., Karaveli, S., Bell, G. R. R., Zia, R., Kuzirian, A. M., Dennis, P. B., Crookes‐Goodson, W. J., Naik, R. R., et al. (2013) · Advanced Functional Materials · doi:10.1002/adfm.201203705 · Cited by 112
The role of protein assembly in dynamically tunable bio-optical tissues
Tao, A. R., DeMartini, D. G., Izumi, M., Sweeney, A., Holt, A. L., & Morse, D. E. (2009) · Biomaterials · doi:10.1016/j.biomaterials.2009.10.038 · Cited by 105
Self-citationPhysiological color change in squid iridophores
Cooper, K. M., Hanlon, R. T., & Budelmann, B. U. (1990) · Cell and Tissue Research · doi:10.1007/bf00571425 · Cited by 102
Physiological color change in squid iridophores
Hanlon, R. T., Cooper, K. M., Budelmann, B. U., & Pappas, T. C. (1990) · Cell and Tissue Research · doi:10.1007/bf00571424 · Cited by 95
Changes in reflectin protein phosphorylation are associated with dynamic iridescence in squid
Izumi, M., Sweeney, A., DeMartini, D. G., Weaver, J. C., Powers, M. L., Tao, A. R., Silvas, T. V., Kramer, R. M., Crookes‐Goodson, W. J., Mäthger, L. M., et al. (2009) · Journal of The Royal Society Interface · doi:10.1098/rsif.2009.0299 · Cited by 85
Self-citationAnatomical basis for camouflaged polarized light communication in squid
Mäthger, L. M., & Hanlon, R. T. (2006) · Biology Letters · doi:10.1098/rsbl.2006.0542 · Cited by 75
Neural control of tuneable skin iridescence in squid
Wardill, T. J., Gonzalez-Bellido, P. T., Crook, R. J., & Hanlon, R. T. (2012) · Proceedings of the Royal Society B Biological Sciences · doi:10.1098/rspb.2012.1374 · Cited by 70
Shashar, N., Vaughan, K., Loew, E. R., Boal, J. G., Hanlon, R. T., & Grable, M. M. (2004) · Behaviour · doi:10.1163/1568539042265662 · Cited by 65
Cephalopod coloration model I Squid chromatophores and iridophores
Sutherland, R. L., Mäthger, L. M., Hanlon, R. T., Urbas, A. M., & Stone, M. O. (2008) · Journal of the Optical Society of America A · doi:10.1364/josaa.25.000588 · Cited by 61
Mäthger, L. M., & Denton, E. J. (2001) · Journal of Experimental Biology · doi:10.1242/jeb.204.12.2103 · Cited by 54
Optical parameters of the tunable Bragg reflectors in squid
Ghoshal, A., DeMartini, D. G., Eck, E., & Morse, D. E. (2013) · Journal of The Royal Society Interface · doi:10.1098/rsif.2013.0386 · Cited by 53
Self-citationChiou, T.-H., Mäthger, L. M., Hanlon, R. T., & Cronin, T. W. (2007) · Journal of Experimental Biology · doi:10.1242/jeb.006932 · Cited by 47
Preliminary notices of some new Pacific cephalopods
Berry, S. S. (1911) · Proceedings of the United States National Museum · doi:10.5479/si.00963801.1838.589 · Cited by 36
Reflector cells in the skin of Octopus dofleini
Brocco, S. L., & Cloney, R. (1980) · Cell and Tissue Research · doi:10.1007/bf00234678 · Cited by 29
Flagged first, then by citations
Research works printed but not linked: 1
- DeMartini D, 2014, Molecular and cellular mechanisms controlling tunable reflectance
Cited by
Sorted by citations
Dynamic pigmentary and structural coloration within cephalopod chromatophore organs
Williams, T. L., Senft, S. L., Yeo, J., Martín-Martínez, F. J., Kuzirian, A. M., Martin, C. A., DiBona, C. W., Chen, C.-T., Dinneen, S. R., Nguyen, H. T., et al. (2019) · Nature Communications · doi:10.1038/s41467-019-08891-x · Cited by 178
Photosymbiotic giant clams are transformers of solar flux
Holt, A. L., Vahidinia, S., Gagnon, Y. L., Morse, D. E., & Sweeney, A. (2014) · Journal of The Royal Society Interface · doi:10.1098/rsif.2014.0678 · Cited by 100
Self-citationLevenson, R., Bracken, C., Bush, N., & Morse, D. E. (2016) · Journal of Biological Chemistry · doi:10.1074/jbc.m115.686014 · Cited by 62
Self-citationCephalopod‐Inspired High Dynamic Range Mechano‐Imaging in Polymeric Materials
Clough, J. M., van der Gucht, J., Kodger, T. E., & Sprakel, J. H. B. (2020) · Advanced Functional Materials · doi:10.1002/adfm.202002716 · Cited by 49
Levenson, R., DeMartini, D. G., & Morse, D. E. (2017) · APL Materials · doi:10.1063/1.4985758 · Cited by 40
Self-citationLevenson, R., Bracken, C. J., Sharma, C., Santos, J., Arata, C., Kohl, P., Li, Y., & Morse, D. E. (2019) · Journal of Biological Chemistry · doi:10.1074/jbc.ra119.010339 · Cited by 41
Self-citationResearch Progress of Bionic Adaptive Camouflage Materials
Qiao, Y., Meng, Z., Wang, P., & Yan, D. (2021) · Frontiers in Materials · doi:10.3389/fmats.2021.637664 · Cited by 38
Cephalopod-inspired optical engineering of human cells
Chatterjee, A., Sanchez, J. A. C., Yamauchi, T., Taupin, V., Couvrette, J., & Gorodetsky, A. A. (2020) · Nature Communications · doi:10.1038/s41467-020-16151-6 · Cited by 37
Wavelength-specific forward scattering of light by Bragg-reflective iridocytes in giant clams
Ghoshal, A., Eck, E., Gordon, M. J., & Morse, D. E. (2016) · Journal of The Royal Society Interface · doi:10.1098/rsif.2016.0285 · Cited by 30
Self-citationAn introduction to color-changing systems from the cephalopod protein reflectin
Chatterjee, A., Norton‐Baker, B., Bagge, L. E., Patel, P., & Gorodetsky, A. A. (2018) · Bioinspiration & Biomimetics · doi:10.1088/1748-3190/aab804 · Cited by 27
Structure, self-assembly, and properties of a truncated reflectin variant
Umerani, M. J., Pratakshya, P., Chatterjee, A., Sanchez, J. A. C., Kim, H. S., Ilc, G., Kovačič, M., Magnan, C., Marmiroli, B., Sartori, B., et al. (2020) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2009044117 · Cited by 25
Tough and Variable-Band-Gap Photonic Hydrogel Displaying Programmable Angle-Dependent Colors
Haque, M. A., Mito, K., Kurokawa, T., Nakajima, T., Nonoyama, T., Ilyas, M., & Gong, J. P. (2018) · ACS Omega · doi:10.1021/acsomega.7b01443 · Cited by 25
Zhang, J., Wang, P., Xie, W., Wang, H., Zhang, Y., & Zhou, H. (2024) · ACS Nano · doi:10.1021/acsnano.4c08338 · Cited by 21
Color Richness in Cephalopod Chromatophores Originating from High Refractive Index Biomolecules
Dinneen, S. R., Osgood, R. M., Greenslade, M. E., & Deravi, L. F. (2016) · The Journal of Physical Chemistry Letters · doi:10.1021/acs.jpclett.6b02398 · Cited by 18
At the Intersection of Natural Structural Coloration and Bioengineering
Chatterjee, A. (2022) · Biomimetics · doi:10.3390/biomimetics7020066 · Cited by 17
Reflectin Proteins Bind and Reorganize Synthetic Phospholipid Vesicles
Song, J., Levenson, R., Santos, J., Velazquez, L. R., Zhang, F., Fygenson, D. K., Wu, W., & Morse, D. E. (2020) · Langmuir · doi:10.1021/acs.langmuir.9b03632 · Cited by 13
Self-citationLoke, J. J., Hoon, S. S., & Miserez, A. (2022) · ACS Applied Materials & Interfaces · doi:10.1021/acsami.2c01999 · Cited by 11
Voltage-calibrated, finely tunable protein assembly
Lin, Y., Masquelier, E., Sabeh, Y. A., Sepunaru, L., Gordon, M. J., & Morse, D. E. (2023) · Journal of The Royal Society Interface · doi:10.1098/rsif.2023.0183 · Cited by 12
Self-citationSquid Skin Cell-Inspired Refractive Index Mapping of Cells, Vesicles, and Nanostructures
Chatterjee, A., Pratakshya, P., Kwansa, A. L., Kaimal, N., Cannon, A. H., Sartori, B., Marmiroli, B., Orins, H., Feng, Z., Drake, S., et al. (2023) · ACS Biomaterials Science & Engineering · doi:10.1021/acsbiomaterials.2c00088 · Cited by 10
Gordon, R., Levenson, R., Malady, B. T., Sabeh, Y. A., Nguyen, A., & Morse, D. E. (2025) · Journal of Biological Chemistry · doi:10.1016/j.jbc.2025.108277 · Cited by 7
Self-citationElectrochemically Driven Optical Dynamics of Reflectin Protein Films
Lin, Y., Yang, C., Tochikura, S. K., Uzarski, J. R., Morse, D. E., Sepunaru, L., & Gordon, M. J. (2025) · Advanced Materials · doi:10.1002/adma.202411005 · Cited by 6
Self-citationDawson, N. J., & Lynch-Holm, V. (2021) · Journal of Luminescence · doi:10.1016/j.jlumin.2021.118474 · Cited by 6
Core–shell nanospheres behind the blue eyes of the bay scallopArgopecten irradians
Harris, O. K., Kingston, A. C. N., Wolfe, C. S., Ghoshroy, S., Johnsen, S., & Speiser, D. I. (2019) · Journal of The Royal Society Interface · doi:10.1098/rsif.2019.0383 · Cited by 5
An iterative correction approach used to retrieve the refractive index of squid pigment aerosols
Dinneen, S. R., Deravi, L. F., & Greenslade, M. E. (2018) · Journal of Optics · doi:10.1088/2040-8986/aaa6ff · Cited by 3
Sakai, D., Nishikawa, J., Kakiuchida, H., & Hirose, E. (2022) · PeerJ · doi:10.7717/peerj.14284 · Cited by 2
FRET Dilution Assay for Analyzing Dynamic Exchange Between Protein Assemblies
Gordon, R., & Morse, D. E. (2025) · Journal of Visualized Experiments · doi:10.3791/68678 · Cited by 1
Self-citationXie, W., Zhang, J., Wang, P., Sun, Y. N., Li, J., Xu, J., Zhang, D., & Zhou, H. (2025) · Nano Letters · doi:10.1021/acs.nanolett.5c02289 · Cited by 1
Elucidating the role of nanostructured pigment granules in the dynamic coloration of cephalopods
Williams, T. L. (2019) · Digital Repository Service (Northeastern University) · doi:10.17760/d20316340
Biomimetic photonic coatings using squid reflectin protein
Loke, J. J. (2022) · DR-NTU (Nanyang Technological University) · doi:10.32657/10356/163521
Dynamic self-assembly of reflectins and their contribution to cephalopods coloration
Soares, C. L., Dias, A. M. G. C., & Roque, A. C. A. (2025) · Biophysics Reviews · doi:10.1063/5.0294035
Gordon, R., Levenson, R., Malady, B. T., Sabeh, Y. A., & Morse, D. E. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.03.13.711711
Self-citationugurlu, S. Y., & Bal, E. (2026) · ChemistrySelect · doi:10.1002/slct.73822
Gordon, R., Levenson, R., Malady, B. T., Sabeh, Y. A., & Morse, D. E. (2026) · Biophysical Journal · doi:10.1016/j.bpj.2026.09.039
Self-citation
Metadata
- Authors
- Ghoshal, A., DeMartini, D. G., Eck, E., & Morse, D. E.
- Affiliations
- University of California, Santa Barbara (US)
- Corresponding author
- Morse, Daniel E.
- Venue
- Journal of The Royal Society Interface
- Publisher
- The Royal Society
- Published
- 2014-06-06
- Volume
- 11
- Issue
- 95
- Pages
- 20140106
- ISSN
- 1742-5662 (ISSN-L)
- 1742-5689
- PMC
- PMC4006249
- OpenAlex
- W2151237885
- Cited by
- 35
- References
- 22
- Citations by year
- 2026: 3 · 2025: 5 · 2024: 1 · 2023: 2 · 2022: 6 · 2021: 2 · 2020: 4 · 2019: 4 · 2018: 3 · 2017: 1 · 2016: 2 · 2015: 1 · 2014: 1
- FWCI
- 0.83
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Journal of The Royal Society Interface (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Topics
- Physical Sciences › Physics and Astronomy › Atomic and Molecular Physics, and Optics › Photonic Crystals and Applications
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Cephalopods and Marine Biology
- Physical Sciences › Materials Science › Biomaterials › Calcium Carbonate Crystallization and Inhibition
- Keywords
- reflectin, refractive index, distributed Bragg reflector, structural color, iridescence, Doryteuthis opalescens, lamellae, camouflage, reflectivity, Mollusca
- MeSH terms
- Animals, Refractometry, Skin, Skin Pigmentation, Decapodiformes
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review
