@article{9b7e428e9d03480bb89199fd70f0826f,
title = "The Making of a Flight Feather: Bio-architectural Principles and Adaptation",
abstract = "The evolution of flight in feathered dinosaurs and early birds over millions of years required flight feathers whose architecture features hierarchical branches. While barb-based feather forms were investigated, feather shafts and vanes are understudied. Here, we take a multi-disciplinary approach to study their molecular control and bio-architectural organizations. In rachidial ridges, epidermal progenitors generate cortex and medullary keratinocytes, guided by Bmp and transforming growth factor β (TGF-β) signaling that convert rachides into adaptable bilayer composite beams. In barb ridges, epidermal progenitors generate cylindrical, plate-, or hooklet-shaped barbule cells that form fluffy branches or pennaceous vanes, mediated by asymmetric cell junction and keratin expression. Transcriptome analyses and functional studies show anterior-posterior Wnt2b signaling within the dermal papilla controls barbule cell fates with spatiotemporal collinearity. Quantitative bio-physical analyses of feathers from birds with different flight characteristics and feathers in Burmese amber reveal how multi-dimensional functionality can be achieved and may inspire future composite material designs. Video Abstract: [Figure presented] The design and developmental paradigms of flight feathers are explored using a combination of bio-physical analyses, molecular characterization, and evolutionary comparisons across a broad range of birds with different flight modes, revealing a modular architectural design that can accommodate diverse eco-spaces.",
keywords = "amber, branching morphogenesis, composite biomaterials, dermal papilla, development, evolution, feathered dinosaurs, keratin, morphogenesis, stem cells",
author = "Chang, {Wei Ling} and Hao Wu and Chiu, {Yu Kun} and Shuo Wang and Jiang, {Ting Xin} and Luo, {Zhong Lai} and Lin, {Yen Cheng} and Ang Li and Hsu, {Jui Ting} and Huang, {Heng Li} and Gu, {How Jen} and Lin, {Tse Yu} and Yang, {Shun Min} and Lee, {Tsung Tse} and Lai, {Yung Chi} and Mingxing Lei and Shie, {Ming You} and Yao, {Cheng Te} and Chen, {Yi Wen} and Tsai, {J. C.} and Shieh, {Shyh Jou} and Hwu, {Yeu Kuang} and Cheng, {Hsu Chen} and Tang, {Pin Chi} and Hung, {Shih Chieh} and Chen, {Chih Feng} and Michael Habib and Widelitz, {Randall B.} and Ping Wu and Juan, {Wen Tau} and Chuong, {Cheng Ming}",
note = "Funding Information: Feather microtome sectioning service was provided by Taiwan Mouse Clinic; high-power x-ray for feather tomography was provided by NSRRC; confocal imaging was performed at core in IBMS and IMB, Academia Sinica, Taiwan, and USC Stem Cell Microscopy Core; and RNA sequencing was performed at Epigenome Core, USC. We thank Hwei-Feng Chang, Hsiang-Hsin Chen, Chia-Ru Chang, Su-Ting Hsu, and Fu-Lang Chan for helping with data acquisition and the staff at LRI, WBST, and NMMBA for providing avian species feather samples. This work is supported by the ISCC, CMUH, Taiwan, the Drug Development Center, CMU, supported by Higher Education Sprout Project by Ministry of Education (HESP-MOE), and grants from Ministry of Science and Technology (MOST 105-2311-B-039-001 and MOST 106-2311-B-039-001), Taiwan. Work at USC is supported by US NIH (AR 047364 and AR 060306). C.-M.C. P.W. T.-X.J. and R.B.W. are partially supported by a research contract between CMUH and USC (USC 5351285884), and C.-M.C. is a paid scientific advisor of CMU. The team at NCHU is supported by iEGG/Avian Genetic Resource/ABC supported by HESP-MOE (107-S-0023). S.W. works in USC via support from the Human Frontier Science Program (LT-000728/2018) and the National Natural Science Foundation of China (NSFC 41602013). Z.-L.L. is a visiting fellow worked at USC via support by NSFC (31370269). Collaborative members are supported by Academia Sinica Research Program on Nanoscience and Nanotechnology; iWRR is supported by Top Notch Project, NCKU (D103-35B05, D104-35B04), and a University Advancement grant by MOE, Taiwan. C.-M.C. dedicates this work to Professor Fon Jou Hsieh for his inspiration. This work is the result of international collaboration by scientific teams from different disciplines. Conceptualization, W.-T.J. W.-L.C. and C.-M.C.; Methodology and Discussion, W.-T.J. W.-L.C. Y.-C. Lai, M.L. H.-C.C. and C.-M.C.; Investigation, W.-L.C. T.-X.J. P.W. Z.-L.L. Y.-C. Lin, S.W. A.L. W.-T.J. H.W. W.-L.C. Y.-K.C. H.-J.G. T.-Y.L. S.-M.Y. M.-Y.S. C.-F.C. P.-C.T. and Y.-W.C.; Formal Analysis, W.-L.C. W.-T.J. H.W. J.-T.H. H.-L.H. T.-Y.L. Y.-K.H. M.H. and P.W.; Writing ? Original Draft, W.-T.J. W.-L.C. J.-T.H. H.-L.H. J.-C.T. Y.-K.H. M.H. P.W. R.B.W. and C.-M.C.; Writing ? Review & Editing, W.-L.C. W.-T.J. P.W. R.B.W. and C.-M.C.; Funding Acquisition, C.-M.C. W.-T.J. S.-C.H. R.B.W. P.W. C.-F.C. P.-C.T. S.-J.S.; Resources, C.-T.Y. Y.-C. Lin, P.-C.T. and C.-F.C.; Supervision, W.-T.J. and C.-M.C. The authors declare no competing interests. Funding Information: Feather microtome sectioning service was provided by Taiwan Mouse Clinic; high-power x-ray for feather tomography was provided by NSRRC; confocal imaging was performed at core in IBMS and IMB, Academia Sinica, Taiwan, and USC Stem Cell Microscopy Core; and RNA sequencing was performed at Epigenome Core, USC. We thank Hwei-Feng Chang, Hsiang-Hsin Chen, Chia-Ru Chang, Su-Ting Hsu, and Fu-Lang Chan for helping with data acquisition and the staff at LRI, WBST, and NMMBA for providing avian species feather samples. This work is supported by the ISCC , CMUH , Taiwan, the Drug Development Center, CMU , supported by Higher Education Sprout Project by Ministry of Education (HESP-MOE), and grants from Ministry of Science and Technology (MOST 105-2311-B-039-001 and MOST 106-2311-B-039-001 ), Taiwan. Work at USC is supported by US NIH ( AR 047364 and AR 060306 ). C.-M.C., P.W., T.-X.J., and R.B.W. are partially supported by a research contract between CMUH and USC (USC 5351285884 ), and C.-M.C. is a paid scientific advisor of CMU. The team at NCHU is supported by iEGG/Avian Genetic Resource/ABC supported by HESP-MOE ( 107-S-0023 ). S.W. works in USC via support from the Human Frontier Science Program ( LT-000728/2018 ) and the National Natural Science Foundation of China (NSFC 41602013 ). Z.-L.L. is a visiting fellow worked at USC via support by NSFC ( 31370269 ). Collaborative members are supported by Academia Sinica Research Program on Nanoscience and Nanotechnology ; iWRR is supported by Top Notch Project , NCKU ( D103-35B05 , D104-35B04 ), and a University Advancement grant by MOE, Taiwan . C.-M.C. dedicates this work to Professor Fon Jou Hsieh for his inspiration. Publisher Copyright: {\textcopyright} 2019 Elsevier Inc.",
year = "2019",
month = nov,
day = "27",
doi = "10.1016/j.cell.2019.11.008",
language = "English",
volume = "179",
pages = "1409--1423.e17",
journal = "Cell",
issn = "0092-8674",
publisher = "Cell Press",
number = "6",
}