{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,7]],"date-time":"2026-10-07T16:08:17Z","timestamp":1791389297228,"version":"4.3.3"},"reference-count":33,"publisher":"Public Library of Science (PLoS)","issue":"8","license":[{"start":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T00:00:00Z","timestamp":1786579200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS One"],"abstract":"<jats:p>\n                    Important in many human artistic cultures, checkerboard patterns are rare in nature like many motifs based on squared geometry. Nevertheless, they are expected to be very detectable by the visual systems due to their periodic geometry and contrasted two-tone coloring, therefore potential specific biological functions are suspected. Here, thanks to a biological survey, we draw the first diversity landscape of eukaryotic species bearing checkerboard patterns, confirming their rarity but also their presence in extremely diverse clades. Then, we selected two genera,\n                    <jats:italic>Sarcophaga<\/jats:italic>\n                    flies and\n                    <jats:italic>Fritillaria<\/jats:italic>\n                    flowers, to perform in-depth pattern analyses allowing us to make strong hypotheses on the mechanisms producing these very peculiar patterns, as no morphogenetic process was known to generate checkerboards. Although they share a similar geometry, these two genera appear to produce checkerboards through very different ways, showing a convergence of shape but not of processes. Whereas the\n                    <jats:italic>Fritillaria<\/jats:italic>\n                    analysis points to a geometric constraining of a Turing-like pattern by the parallel network of veins, that of\n                    <jats:italic>Sarcophaga<\/jats:italic>\n                    suggest the reuse of developmental boundaries and right-left symmetry, together by the combination of vertical and horizontal stripes. Furthermore, we present the first description to our knowledge of the striking color-changing nature of\n                    <jats:italic>Sarcophaga<\/jats:italic>\n                    checkerboards, whose light and dark squares can exchange their color depending on the angles of lighting and observation thanks to the planar polarity of the cuticular hairs, the setae. Together, this shows the extent of the processes selected during evolution to generate complex forms and colors, and confirms the importance of studying morphogenesis with in-depth pattern analyses and through species diversity. Finally, by enabling strong hypotheses to be made about the morphogenesis of these patterns, it paves the way for the molecular identification of the morphogenetic processes at work.\n                  <\/jats:p>","DOI":"10.1371\/journal.pone.0352968","type":"journal-article","created":{"date-parts":[[2026,8,13]],"date-time":"2026-08-13T17:40:18Z","timestamp":1786642818000},"page":"e0352968","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":1,"title":["Changing rounds into squares or combining stripes? In-depth analysis of Fritillaria flowers and the color-changing pattern of Sarcophaga flies sheds light on the diversity and morphogenesis of checkerboard patterns in Eukaryotes"],"prefix":"10.1371","volume":"21","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-2335-9556","authenticated-orcid":true,"given":"Pierre","family":"Galipot","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Julie","family":"Zalko","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laetitia","family":"Carrive","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Romain","family":"Garrouste","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Agn\u00e8s","family":"Schermann-Legionnet","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,8,13]]},"reference":[{"issue":"3","key":"pone.0352968.ref001","doi-asserted-by":"crossref","first-page":"037114","DOI":"10.1063\/1.2214167","article-title":"Turing patterns beyond hexagons and stripes","volume":"16","author":"L Yang","year":"2006","journal-title":"Chaos"},{"issue":"2","key":"pone.0352968.ref002","doi-asserted-by":"crossref","first-page":"023115","DOI":"10.1063\/1.4875262","article-title":"Square Turing patterns in reaction-diffusion systems with coupled layers","volume":"24","author":"J Li","year":"2014","journal-title":"Chaos"},{"issue":"6046","key":"pone.0352968.ref003","doi-asserted-by":"crossref","first-page":"1144","DOI":"10.1126\/science.1208467","article-title":"Nectins establish a checkerboard-like cellular pattern in the auditory epithelium","volume":"333","author":"H Togashi","year":"2011","journal-title":"Science"},{"issue":"3","key":"pone.0352968.ref004","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1387\/ijdb.1702680","article-title":"A checkerboard pattern manifested by the oviduct epithelium of the Japanese quail","volume":"34","author":"HI Yamanaka","year":"1990","journal-title":"Int J Dev Biol"},{"issue":"21","key":"pone.0352968.ref005","doi-asserted-by":"crossref","DOI":"10.1242\/dev.201519","article-title":"The shift to 3D growth during embryogenesis of kelp species, atlas of cell division and differentiation of Saccharina latissima","volume":"150","author":"I Theodorou","year":"2023","journal-title":"Development"},{"key":"pone.0352968.ref006","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1113\/jphysiol.1979.sp012827","article-title":"Responses of striate cortex cells to grating and checkerboard patterns","volume":"291","author":"KK De Valois","year":"1979","journal-title":"J Physiol"},{"issue":"4","key":"pone.0352968.ref007","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1167\/jov.22.4.3","article-title":"Mapping spatial frequency preferences across human primary visual cortex","volume":"22","author":"WF Broderick","year":"2022","journal-title":"J Vis"},{"issue":"22","key":"pone.0352968.ref008","doi-asserted-by":"crossref","first-page":"8504","DOI":"10.1523\/JNEUROSCI.20-22-08504.2000","article-title":"Spatial frequency maps in cat visual cortex","volume":"20","author":"NP Issa","year":"2000","journal-title":"J Neurosci"},{"issue":"5","key":"pone.0352968.ref009","doi-asserted-by":"crossref","first-page":"41","DOI":"10.1007\/s11538-021-00870-y","article-title":"Bespoke Turing Systems","volume":"83","author":"TE Woolley","year":"2021","journal-title":"Bull Math Biol"},{"issue":"5","key":"pone.0352968.ref010","first-page":"133","article-title":"A monograph of the North American species of Fritillaria","volume":"7","author":"DE Beetle","year":"1944","journal-title":"Madro\u00f1o"},{"key":"pone.0352968.ref011","doi-asserted-by":"crossref","DOI":"10.1017\/qpb.2025.8","article-title":"Colour pattern studies: the SE (\u305b) method, a shape-centred approach to explore biodiversity and avoid aesthetic biases","volume":"6","author":"P Galipot","year":"2025","journal-title":"Quant Plant Biol"},{"issue":"2","key":"pone.0352968.ref012","doi-asserted-by":"crossref","first-page":"105","DOI":"10.1007\/s13752-024-00477-1","article-title":"Biological theories of morphogenesis based on holistic biophysical thinking","volume":"20","author":"KH Palmquist","year":"2025","journal-title":"Biol Theory"},{"key":"pone.0352968.ref013","year":"2025"},{"issue":"2","key":"pone.0352968.ref014","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0305921","article-title":"And growth on form? 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