{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,19]],"date-time":"2026-08-19T19:30:08Z","timestamp":1787167808835,"version":"build-2736575974"},"reference-count":74,"publisher":"Wiley","issue":"16","license":[{"start":{"date-parts":[[2026,4,2]],"date-time":"2026-04-02T00:00:00Z","timestamp":1775088000000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,4,2]],"date-time":"2026-04-02T00:00:00Z","timestamp":1775088000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32400988"],"award-info":[{"award-number":["32400988"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100015401","name":"Key Research and Development Projects of Shaanxi Province","doi-asserted-by":"publisher","award":["2022SF\u2010276"],"award-info":[{"award-number":["2022SF\u2010276"]}],"id":[{"id":"10.13039\/501100015401","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["febs.onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["The FEBS Journal"],"published-print":{"date-parts":[[2026,8]]},"abstract":"<jats:p>\n                    Animal body color patterning, as a naturally selected adaptive trait, plays critical ecological roles (e.g., warning coloration and mimicry). These patterns develop through spatiotemporally precise effector gene expression (e.g., pigment synthetases), regulated by pattern genes. However, the key pattern genes and the underlying mechanisms that govern the formation of marking patterns remain poorly defined. This study investigated the silkworm mutant, multilunar (\n                    <jats:italic>L<\/jats:italic>\n                    ), which exhibited twin\u2010spot markings on sequential segments. We found that\n                    <jats:italic>Wnt6<\/jats:italic>\n                    mRNA expression was significantly upregulated in the twin\u2010spot epidermal region. The\n                    <jats:italic>cuticular protein RR\u20101 motif 4<\/jats:italic>\n                    (\n                    <jats:italic>CPR4<\/jats:italic>\n                    ) and\n                    <jats:italic>cuticular protein RR\u20101 motif 38<\/jats:italic>\n                    (\n                    <jats:italic>CPR38<\/jats:italic>\n                    ) promoters were identified as core cis\u2010acting elements for epidermal specificity throughout larval development.\n                    <jats:italic>CPR4<\/jats:italic>\n                    \u2010driven\n                    <jats:italic>Wnt6<\/jats:italic>\n                    overexpression induced twin\u2010spot markings on the epidermis of silkworm larvae, resembling the mutant\n                    <jats:italic>L<\/jats:italic>\n                    pattern, while exhibiting growth advantages compared with the\n                    <jats:italic>CPR38<\/jats:italic>\n                    \u2010driven system. Ultrastructural analysis indicated that there was a positive correlation between epidermal bulge structures and the formation of twin spots. Mechanistically, quantitative real\u2010time PCR (qRT\u2010PCR) analysis confirmed that epidermis\u2010specific overexpression of\n                    <jats:italic>Wnt6<\/jats:italic>\n                    did not induce co\u2010activation of coclustered Wnt members. Furthermore, the analysis results of RNA\u2010seq and qRT\u2010PCR suggest that\n                    <jats:italic>Wnt6<\/jats:italic>\n                    may promote the formation of the twin\u2010spot patterns by participating in hormone regulation, pigment production, and cell proliferation signaling pathways. This study, for the first time, establishes a heritable\n                    <jats:italic>Wnt6<\/jats:italic>\n                    ectopic\u2010expression system that induces a twin\u2010spot phenotype resembling that of the mutant\n                    <jats:italic>L<\/jats:italic>\n                    , thereby providing a potential molecular target for elucidating the evolutionary mechanisms of adaptive body coloration in insects.\n                  <\/jats:p>","DOI":"10.1111\/febs.70491","type":"journal-article","created":{"date-parts":[[2026,4,3]],"date-time":"2026-04-03T05:11:17Z","timestamp":1775193077000},"page":"4916-4942","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Ectopic\n                    <i>Wnt6<\/i>\n                    expression induces twin\u2014spot markings in the epidermis of the silkworm\n                    <i>Bombyx mori<\/i>\n                    \u2013 Potential molecular target to discover evolutionary mechanisms of adaptive body coloration in insects"],"prefix":"10.1111","volume":"293","author":[{"ORCID":"https:\/\/orcid.org\/0009-0000-3581-9217","authenticated-orcid":false,"given":"Xuan","family":"Xie","sequence":"first","affiliation":[{"name":"The Key Laboratory of Aerospace Medicine, Ministry of Education Fourth Military Medical University  Xi'an China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhijun","family":"Yan","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Aerospace Medicine, Ministry of Education Fourth Military Medical University  Xi'an China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ziyu","family":"Jin","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Aerospace Medicine, Ministry of Education Fourth Military Medical University  Xi'an China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yijin","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Life Sciences Northwest University  Xi'an China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xian","family":"Guo","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Aerospace Medicine, Ministry of Education Fourth Military Medical University  Xi'an China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-9387-9058","authenticated-orcid":false,"given":"Jiangdong","family":"Zhao","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Aerospace Medicine, Ministry of Education Fourth Military Medical University  Xi'an China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9257-206X","authenticated-orcid":false,"given":"Fan","family":"Jiang","sequence":"additional","affiliation":[{"name":"Section of Health No. 94804 Unit of the Chinese People's Liberation Army  Shanghai China"},{"name":"Resident Standardization Training Cadet Corps Air Force Medical Center  Beijing China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7463-8379","authenticated-orcid":false,"given":"Ru","family":"Zhang","sequence":"additional","affiliation":[{"name":"The Key Laboratory of Aerospace Medicine, Ministry of Education Fourth Military Medical University  Xi'an China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,4,2]]},"reference":[{"key":"e_1_2_10_2_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41576-020-0234-z"},{"key":"e_1_2_10_3_1","first-page":"1277","article-title":"Studies on the mutant systems of the Bombyx mori gene bank","volume":"1","author":"Lu C","year":"2022","journal-title":"Agric Sci 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