{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,19]],"date-time":"2026-06-19T07:52:10Z","timestamp":1781855530878,"version":"3.54.5"},"reference-count":48,"publisher":"Wiley","license":[{"start":{"date-parts":[[2026,6,19]],"date-time":"2026-06-19T00:00:00Z","timestamp":1781827200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/onlinelibrary.wiley.com\/termsAndConditions#vor"},{"start":{"date-parts":[[2026,6,19]],"date-time":"2026-06-19T00:00:00Z","timestamp":1781827200000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100007540","name":"Jiangsu Agricultural Science and Technology Innovation Fund","doi-asserted-by":"publisher","award":["CX(20)2029"],"award-info":[{"award-number":["CX(20)2029"]}],"id":[{"id":"10.13039\/100007540","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Insect Science"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    The silkworm,\n                    <jats:italic>Bombyx mori<\/jats:italic>\n                    , is an economically important insect resource in China. With the transformation and upgrading of the sericulture industry, genetically engineered silkworms have emerged as efficient animal bioreactors for producing bioactive proteins and functional compounds. However, the diversity of available transgenic silkworm bioreactor resources remains limited and requires further expansion. Herein, we focused on human lactoferrin (hLF), which exhibits potent antibacterial activity due to its strong iron\u2010chelating capacity, and established an efficient recombinant hLF (rhLF) expression system in the posterior silk glands (PSGs) of transgenic silkworms using\n                    <jats:italic>piggyBac<\/jats:italic>\n                    technology. The rhLF production reached 45.13 \u00b1 2.46 mg\/g cocoon shell weight, with N\u2010glycosylation modifications consistent with those of native hLF. Functionally, rhLF exhibited robust antibacterial activity, achieving inhibition rates of 31.64% \u00b1 4.66% against\n                    <jats:italic>Escherichia coli<\/jats:italic>\n                    and 74.35% \u00b1 0.87% against\n                    <jats:italic>Staphylococcus aureus<\/jats:italic>\n                    . Additionally, rhLF significantly promoted cell migration and accelerated wound healing, with a migration rate of 17.06% \u00b1 4.05%. The resulting chimeric fibroin fibers containing rhLF demonstrated enhanced antibacterial activity while maintaining cytocompatibility. Collectively, we constructed a novel transgenic silkworm strain enabling high\u2010yield rhLF production and achieved the biomanufacturing of functional silk fibers. These findings provide a cost\u2010effective strategy for generating high\u2010value products within conventional agricultural systems.\n                  <\/jats:p>","DOI":"10.1111\/1744-7917.70317","type":"journal-article","created":{"date-parts":[[2026,6,19]],"date-time":"2026-06-19T07:13:56Z","timestamp":1781853236000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A genetically engineered silkworm bioreactor enabling efficient recombinant human lactoferrin production for functional silk fiber biomanufacturing"],"prefix":"10.1111","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5953-3624","authenticated-orcid":false,"given":"Yi\u2010Jiangcheng","family":"Li","sequence":"first","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shi\u2010Hua","family":"Li","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Da\u2010Lin","family":"Zhu","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-7552-1862","authenticated-orcid":false,"given":"Yu\u2010Tong","family":"Liu","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shan\u2010Shan","family":"Wang","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kai","family":"Chen","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"},{"name":"Key Laboratory of Silkworm and Mulberry Genetic Improvement, Ministry of Agriculture and Rural Affairs, Sericultural Scientific Research Center Chinese Academy of Agricultural Sciences Zhenjiang Jiangsu Province China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2764-8125","authenticated-orcid":false,"given":"Sheng","family":"Sheng","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"},{"name":"Key Laboratory of Silkworm and Mulberry Genetic Improvement, Ministry of Agriculture and Rural Affairs, Sericultural Scientific Research Center Chinese Academy of Agricultural Sciences Zhenjiang Jiangsu Province China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2569-1461","authenticated-orcid":false,"given":"Jun","family":"Wang","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"},{"name":"Key Laboratory of Silkworm and Mulberry Genetic Improvement, Ministry of Agriculture and Rural Affairs, Sericultural Scientific Research Center Chinese Academy of Agricultural Sciences Zhenjiang Jiangsu Province China"},{"name":"Jiangsu Key Laboratory of Clean Energy Storage and Conversion, School of Resources and Environmental Engineering Jiangsu University of Technology  Changzhou Jiangsu Province China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"An\u2010Jiang","family":"Tan","sequence":"additional","affiliation":[{"name":"Jiangsu Key Laboratory of Sericultural and Animal Biotechnology, School of Biotechnology Jiangsu University of Science and Technology  Zhenjiang China"},{"name":"Key Laboratory of Silkworm and Mulberry Genetic Improvement, Ministry of Agriculture and Rural Affairs, Sericultural Scientific Research Center Chinese Academy of Agricultural Sciences Zhenjiang Jiangsu Province China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,6,19]]},"reference":[{"key":"e_1_2_7_2_1","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1111\/phen.12421","article-title":"Silkworm Bombyx mori as a model organism: a review","volume":"48","author":"Ashraf H.","year":"2023","journal-title":"Physiological Entomology"},{"key":"e_1_2_7_3_1","doi-asserted-by":"crossref","first-page":"236","DOI":"10.1038\/nature08962","article-title":"Self\u2010assembly of spider silk proteins is controlled by a pH\u2010sensitive relay","volume":"465","author":"Askarieh G.","year":"2010","journal-title":"Nature"},{"key":"e_1_2_7_4_1","doi-asserted-by":"crossref","DOI":"10.3389\/fnut.2022.1018336","article-title":"Lactoferrin: a glycoprotein that plays an active role in human health","volume":"9","author":"Cao X.","year":"2023","journal-title":"Frontiers in Nutrition"},{"key":"e_1_2_7_5_1","doi-asserted-by":"crossref","DOI":"10.1073\/pnas.2306322120","article-title":"Engineering a complex, multiple enzyme\u2010mediated synthesis of natural plant pigments in the silkworm, Bombyx mori","volume":"120","author":"Chen K.","year":"2023","journal-title":"Proceedings of the National Academy of Sciences USA"},{"key":"e_1_2_7_6_1","doi-asserted-by":"crossref","first-page":"2612","DOI":"10.1002\/jbm.b.33804","article-title":"Antimicrobial activity of immobilized lactoferrin and lactoferricin","volume":"105","author":"Chen R.X.","year":"2017","journal-title":"Journal of Biomedical Materials Research. Part B, Applied Biomaterials"},{"key":"e_1_2_7_7_1","doi-asserted-by":"crossref","first-page":"565","DOI":"10.1016\/S0092-8674(04)00130-8","article-title":"Structure of the human transferrin receptor\u2010transferrin complex","volume":"116","author":"Cheng Y.F.","year":"2004","journal-title":"Cell"},{"key":"e_1_2_7_8_1","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1016\/j.bioactmat.2025.06.017","article-title":"Biosynthesis of a dual growth factors (GFs) functionalized silk sericin hydrogel to promote chronic wound healing in diabetic mice","volume":"52","author":"Deng H.","year":"2025","journal-title":"Bioactive Materials"},{"key":"e_1_2_7_9_1","doi-asserted-by":"crossref","first-page":"28","DOI":"10.1016\/j.addr.2019.09.003","article-title":"Silk fibroin for skin injury repair: where do things stand?","volume":"153","author":"Gholipourmalekabadi M.","year":"2020","journal-title":"Advanced Drug Delivery 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Nanobiotechnology"},{"key":"e_1_2_7_13_1","doi-asserted-by":"crossref","first-page":"1445","DOI":"10.20964\/2017.02.57","article-title":"Electrochemical studies on bactericidal mechanism of human lactoferrin and its demand for microenvironment","volume":"12","author":"Huang J.Y.","year":"2017","journal-title":"International Journal of Electrochemical Science"},{"key":"e_1_2_7_14_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.ijbiomac.2023.123863","article-title":"Genetically engineered blue silkworm capable of synthesizing natural blue pigment","volume":"235","author":"Jia L.","year":"2023","journal-title":"International Journal of Biological Macromolecules"},{"key":"e_1_2_7_15_1","doi-asserted-by":"crossref","first-page":"345","DOI":"10.1007\/s43393-021-00066-4","article-title":"Heterologous expression of bovine lactoferrin C\u2010lobe in Bacillus subtilis and comparison of its antibacterial activity with N\u2010lobe","volume":"2","author":"Jin L.","year":"2022","journal-title":"Systems Microbiology and Biomanufacturing"},{"key":"e_1_2_7_16_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.ibmb.2024.104177","article-title":"Binding characteristics and structural dynamics of two general odorant\u2010binding proteins with plant volatiles in the olfactory recognition of Glyphodes pyloalis","volume":"173","author":"Li Y.","year":"2024","journal-title":"Insect Biochemistry and Molecular Biology"},{"key":"e_1_2_7_17_1","doi-asserted-by":"crossref","first-page":"151","DOI":"10.1515\/BC.2001.023","article-title":"Glycoproteins from insect cells: sialylated or not?","volume":"382","author":"Marchal I.","year":"2001","journal-title":"Biological Chemistry"},{"key":"e_1_2_7_18_1","doi-asserted-by":"crossref","first-page":"22","DOI":"10.1139\/bcb-2016-0066","article-title":"Lactoferrin and prematurity: a promising milk protein?","volume":"95","author":"Ochoa T.J.","year":"2017","journal-title":"Biochemistry and Cell 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Biotechnology"},{"key":"e_1_2_7_25_1","doi-asserted-by":"crossref","first-page":"6766","DOI":"10.1073\/pnas.1221700110","article-title":"Transgene\u2010based, female\u2010specific lethality system for genetic sexing of the silkworm, Bombyx mori","volume":"110","author":"Tan A.","year":"2013","journal-title":"Proceedings of the National Academy of Sciences USA"},{"key":"e_1_2_7_26_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.ijbiomac.2023.125527","article-title":"Large\u2010scale and cost\u2010effective production of recombinant human serum albumin (rHSA) in transgenic Bombyx mori cocoons","volume":"245","author":"Tan H.","year":"2023","journal-title":"International Journal of Biological Macromolecules"},{"key":"e_1_2_7_27_1","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1007\/s10529-010-0498-z","article-title":"Transgenic silkworms that weave recombinant proteins into silk cocoons","volume":"33","author":"Tomita M.","year":"2011","journal-title":"Biotechnology Letters"},{"key":"e_1_2_7_28_1","doi-asserted-by":"crossref","first-page":"875","DOI":"10.1248\/yakushi.17-00202-2","article-title":"Development of large scale silkworm\u2010rearing technologies for the GMP production of biologics","volume":"138","author":"Tomita M.","year":"2018","journal-title":"Yakugaku Zasshi\u2010Journal of the Pharmaceutical Society of Japan"},{"key":"e_1_2_7_29_1","doi-asserted-by":"crossref","first-page":"249","DOI":"10.1007\/s10534-006-9062-7","article-title":"A structural comparison of human serum transferrin and human lactoferrin","volume":"20","author":"Wally J.","year":"2007","journal-title":"Biometals"},{"key":"e_1_2_7_30_1","doi-asserted-by":"crossref","first-page":"580","DOI":"10.1080\/10408398.2017.1381583","article-title":"Lactoferrin: structure, function, denaturation and digestion","volume":"59","author":"Wang B.","year":"2019","journal-title":"Critical Reviews in Food Science and 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F.","year":"2024","journal-title":"Insect Science"},{"key":"e_1_2_7_34_1","doi-asserted-by":"crossref","first-page":"925","DOI":"10.1007\/s11248-013-9695-6","article-title":"An optimized sericin\u20101 expression system for mass\u2010producing recombinant proteins in the middle silk glands of transgenic silkworms","volume":"22","author":"Wang F.","year":"2013","journal-title":"Transgenic Research"},{"key":"e_1_2_7_35_1","doi-asserted-by":"crossref","first-page":"1203","DOI":"10.1016\/j.bbrc.2006.01.081","article-title":"In vivo analysis of fibroin heavy chain signal peptide of silkworm Bombyx mori using recombinant baculovirus as vector","volume":"341","author":"Wang S.P.","year":"2006","journal-title":"Biochemical and Biophysical Research Communications"},{"key":"e_1_2_7_36_1","doi-asserted-by":"crossref","first-page":"507","DOI":"10.1111\/j.1745-7270.2006.00189.x","article-title":"Structural analysis of fibroin heavy chain signal peptide of silkworm Bombyx mori","volume":"38","author":"Wang S.P.","year":"2006","journal-title":"Acta Biochimica et Biophysica Sinica"},{"key":"e_1_2_7_37_1","doi-asserted-by":"crossref","first-page":"8757","DOI":"10.1073\/pnas.1806805115","article-title":"Mass spider silk production through targeted gene replacement in Bombyx mori","volume":"115","author":"Xu J.","year":"2018","journal-title":"Proceedings of the National Academy of Sciences USA"},{"key":"e_1_2_7_38_1","doi-asserted-by":"crossref","first-page":"61","DOI":"10.1186\/s13036-019-0186-z","article-title":"A silkworm based silk gland bioreactor for high\u2010efficiency production of recombinant human lactoferrin with antibacterial and anti\u2010inflammatory activities","volume":"13","author":"Xu S.","year":"2019","journal-title":"Journal of Biological Engineering"},{"key":"e_1_2_7_39_1","doi-asserted-by":"crossref","first-page":"6559","DOI":"10.1021\/jf300794h","article-title":"Lowering the blood glucose of diabetes mellitus mice by oral administration with transgenic human insulin\u2010like growth factor I silkworms","volume":"60","author":"Xue R.","year":"2012","journal-title":"Journal of Agricultural and Food Chemistry"},{"key":"e_1_2_7_40_1","doi-asserted-by":"crossref","first-page":"289","DOI":"10.1016\/j.actbio.2011.09.037","article-title":"Macro\/microporous silk fibroin scaffolds with potential for articular cartilage and meniscus tissue engineering applications","volume":"8","author":"Yan L.P.","year":"2012","journal-title":"Acta Biomaterialia"},{"key":"e_1_2_7_41_1","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1016\/j.biotechadv.2017.11.002","article-title":"Engineering strategies for enhanced production of protein and bio\u2010products in Pichia pastoris: a review","volume":"36","author":"Yang Z.L.","year":"2018","journal-title":"Biotechnology Advances"},{"key":"e_1_2_7_42_1","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1093\/jb\/mvm187","article-title":"Effect of N\u2010terminal residues on the structural stability of recombinant horse L\u2010chain apoferritin in an acidic environment","volume":"142","author":"Yoshizawa K.","year":"2007","journal-title":"Journal of Biochemistry"},{"key":"e_1_2_7_43_1","doi-asserted-by":"crossref","first-page":"653","DOI":"10.1007\/s11248-017-0034-1","article-title":"Transgenic silkworms secrete the recombinant glycosylated MRJP1 protein of Chinese honeybee, Apis cerana cerana","volume":"26","author":"You Z.","year":"2017","journal-title":"Transgenic Research"},{"key":"e_1_2_7_44_1","doi-asserted-by":"crossref","first-page":"206","DOI":"10.1093\/glycob\/cwq151","article-title":"Comprehensive characterization of the site\u2010specific N\u2010glycosylation of wild\u2010type and recombinant human lactoferrin expressed in the milk of transgenic cloned cattle","volume":"21","author":"Yu T.","year":"2011","journal-title":"Glycobiology"},{"key":"e_1_2_7_45_1","doi-asserted-by":"crossref","first-page":"128","DOI":"10.1093\/pnasnexus\/pgae128","article-title":"Custom\u2010designed, mass silk production in genetically engineered silkworms","volume":"3","author":"Yu Y.","year":"2024","journal-title":"PNAS Nexus"},{"key":"e_1_2_7_46_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.jinsphys.2021.104265","article-title":"Production of cloned transgenic silkworms by breeding non\u2010diapausing parthenogenetic strains","volume":"132","author":"Zabelina V.","year":"2021","journal-title":"Journal of Insect Physiology"},{"key":"e_1_2_7_47_1","doi-asserted-by":"crossref","DOI":"10.1016\/j.ijbiomac.2023.127766","article-title":"Site\u2010specific glycoproteomic analysis of purified lactoferrin from human and animal milk","volume":"254","author":"Zhang Y.Q.","year":"2024","journal-title":"International Journal of Biological Macromolecules"},{"key":"e_1_2_7_48_1","doi-asserted-by":"crossref","first-page":"29","DOI":"10.1007\/s11248-009-9295-7","article-title":"New and highly efficient expression systems for expressing selectively foreign protein in the silk glands of transgenic silkworm","volume":"19","author":"Zhao A.","year":"2010","journal-title":"Transgenic Research"},{"key":"e_1_2_7_49_1","doi-asserted-by":"crossref","first-page":"216","DOI":"10.1016\/j.actbio.2025.07.030","article-title":"Genetically engineered sericin hydrogels for the delivery of human adiponectin in treating ulcerative colitis in mice","volume":"202","author":"Zhou H.","year":"2025","journal-title":"Acta Biomaterialia"}],"container-title":["Insect 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