{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T05:19:13Z","timestamp":1787635153685,"version":"build-2736575974"},"reference-count":41,"publisher":"MDPI AG","issue":"9","license":[{"start":{"date-parts":[[2026,8,23]],"date-time":"2026-08-23T00:00:00Z","timestamp":1787443200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"publisher","award":["32372943; 32172792"],"award-info":[{"award-number":["32372943; 32172792"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"award":["32372943; 32172792"],"award-info":[{"award-number":["32372943; 32172792"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]},{"name":"China Agriculture Research System","award":["CARS-44-KXJ7"],"award-info":[{"award-number":["CARS-44-KXJ7"]}]},{"award":["CARS-44-KXJ7"],"award-info":[{"award-number":["CARS-44-KXJ7"]}],"id":[{"id":"https:\/\/ror.org\/03p5eky76","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100003392","name":"Natural Science Foundation of Fujian Province","doi-asserted-by":"publisher","award":["2025J01616"],"award-info":[{"award-number":["2025J01616"]}],"id":[{"id":"10.13039\/501100003392","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Biomolecules"],"abstract":"<jats:p>Chalkbrood disease caused by Ascosphaera apis threatens honey bee brood, yet the transcriptional mechanisms that coordinate antifungal immune responses in Apis cerana remain unclear. Here, RNA interference (RNAi) was used to examine two Dorsal homologs and selected dorsal1 transcript isoforms during larval infection. A. apis inoculation increased dorsal1, dorsal2, and five antimicrobial peptide (AMP) transcripts in larval midguts, consistent with an infection-associated humoral response. Gene-level RNAi produced selective transcript responses: dorsal1 knockdown was accompanied by reduced apidaecin and defensin1 expression, whereas dorsal2 knockdown was accompanied principally by reduced defensin1 expression. abaecin, defensin2, and hymenoptaecin transcripts were not significantly altered after dorsal knockdown under the tested conditions. Isoform-targeted RNAi of RNA9886, RNA9888, and RNA9890 was likewise associated with distinct AMP transcript responses, with RNA9888 and RNA9890 more closely associated with defensin1. These transcript-level data support selective and partially overlapping Dorsal-associated regulation during the A. cerana larval response to A. apis, while direct differences in Dorsal protein abundance or activity remain to be established.<\/jats:p>","DOI":"10.3390\/biom16091223","type":"journal-article","created":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T08:03:41Z","timestamp":1787558621000},"page":"1223","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Homologs and Transcriptional Isoforms of Dorsal Are Involved in the Response of Apis cerana to Ascosphaera apis Infection"],"prefix":"10.3390","volume":"16","author":[{"given":"He","family":"Zang","sequence":"first","affiliation":[{"name":"College of Animal Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"},{"name":"College of Bee Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"},{"name":"National & Local United Engineering Laboratory of Natural Biotoxin, Fuzhou 350002, China"},{"name":"Apitherapy Research Institute of Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xinrui","family":"Chen","sequence":"additional","affiliation":[{"name":"College of Bee Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiang","family":"Li","sequence":"additional","affiliation":[{"name":"College of Bee Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xue","family":"Yang","sequence":"additional","affiliation":[{"name":"College of Bee Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qingwei","family":"Tan","sequence":"additional","affiliation":[{"name":"College of Bee Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"},{"name":"National & Local United Engineering Laboratory of Natural Biotoxin, Fuzhou 350002, China"},{"name":"Apitherapy Research Institute of Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dafu","family":"Chen","sequence":"additional","affiliation":[{"name":"College of Bee Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"},{"name":"National & Local United Engineering Laboratory of Natural Biotoxin, Fuzhou 350002, China"},{"name":"Apitherapy Research Institute of Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rui","family":"Guo","sequence":"additional","affiliation":[{"name":"College of Bee Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"},{"name":"National & Local United Engineering Laboratory of Natural Biotoxin, Fuzhou 350002, China"},{"name":"Apitherapy Research Institute of Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4262-9013","authenticated-orcid":false,"given":"Jianfeng","family":"Qiu","sequence":"additional","affiliation":[{"name":"College of Bee Science, Fujian Agriculture and Forestry University, Fuzhou 350002, China"},{"name":"National & Local United Engineering Laboratory of Natural Biotoxin, Fuzhou 350002, China"},{"name":"Apitherapy Research Institute of Fujian Agriculture and Forestry University, Fuzhou 350002, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,23]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Calderone, N.W. (2012). Insect pollinated crops, insect pollinators and US agriculture: Trend analysis of aggregate data for the period 1992\u20132009. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0037235"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"S20","DOI":"10.1016\/j.jip.2009.06.018","article-title":"Chalkbrood disease in honey bees","volume":"103","author":"Aronstein","year":"2010","journal-title":"J. Invertebr. Pathol."},{"key":"ref_3","first-page":"539","article-title":"Analysis of the differentially expressed genes in the 6-day-old larval gut of Apis cerana cerana under the stress of Ascosphaera apis","volume":"39","author":"Guo","year":"2017","journal-title":"J. Environ. Entomol."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2656530","DOI":"10.1080\/21505594.2026.2656530","article-title":"Novel_circ_002651 regulates the immune defense of eastern honeybee larvae against fungal invasion through sponging miR-6001-y","volume":"17","author":"Zhang","year":"2026","journal-title":"Virulence"},{"key":"ref_5","first-page":"12","article-title":"Infection and immunity in the honey bee Apis mellifera","volume":"36","author":"Jarosz","year":"2001","journal-title":"Apiacta"},{"key":"ref_6","unstructured":"Maramorosch, K., and Shope, R.E. (1975). The invertebrate gut as a barrier to invading parasites. Invertebrate Immunity: Mechanisms of Invertebrate Vector\u2013Parasite Relations, Academic Press."},{"key":"ref_7","first-page":"183","article-title":"Response of the Apoidea to fungal infections","volume":"38","author":"Buczek","year":"2003","journal-title":"Apiacta"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"21121","DOI":"10.1074\/jbc.M313856200","article-title":"Differential activation of the NF-\u03baB-like factors Relish and Dif in Drosophila melanogaster by fungi and Gram-positive bacteria","volume":"279","author":"Olcott","year":"2004","journal-title":"J. Biol. Chem."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1159\/000210371","article-title":"Eicosanoid actions in insect immunity","volume":"1","author":"Stanley","year":"2009","journal-title":"J. Innate Immun."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"28569","DOI":"10.1016\/S0021-9258(19)61943-5","article-title":"Acute transcriptional response of the honeybee peptide-antibiotics gene repertoire and required post-translational conversion of the precursor structures","volume":"269","author":"Zhang","year":"1994","journal-title":"J. Biol. Chem."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Zhou, L., Meng, G., Zhu, L., Ma, L., and Chen, K. (2024). Insect antimicrobial peptides as guardians of immunity and beyond: A review. Int. J. Mol. Sci., 25.","DOI":"10.20944\/preprints202403.0123.v1"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"2924","DOI":"10.1038\/s41467-026-69073-0","article-title":"Gut microbiota-derived butyrate primes systemic immunity in honey bees by mediating lipid metabolic reprogramming","volume":"17","author":"Liu","year":"2026","journal-title":"Nat. Commun."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"973","DOI":"10.1016\/S0092-8674(00)80172-5","article-title":"The dorsoventral regulatory gene cassette sp\u00e4tzle\/Toll\/cactus controls the potent antifungal response in Drosophila adults","volume":"86","author":"Lemaitre","year":"1996","journal-title":"Cell"},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"14614","DOI":"10.1073\/pnas.94.26.14614","article-title":"Drosophila host defense: Differential induction of antimicrobial peptide genes after infection by various classes of microorganisms","volume":"94","author":"Lemaitre","year":"1997","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"393","DOI":"10.1146\/annurev.cellbio.12.1.393","article-title":"A conserved signaling pathway: The Drosophila Toll\u2013Dorsal pathway","volume":"12","author":"Belvin","year":"1996","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"645","DOI":"10.1111\/j.1365-2583.2006.00682.x","article-title":"Immune pathways and defence mechanisms in honey bees Apis mellifera","volume":"15","author":"Evans","year":"2006","journal-title":"Insect Mol. Biol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1146\/annurev-cellbio-120319-120223","article-title":"Neofunctionalization of Toll signaling in insects: From immunity to dorsoventral patterning","volume":"39","author":"Roth","year":"2023","journal-title":"Annu. Rev. Cell Dev. Biol."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Liu, G., Tian, Y., Hanson, M.A., Sah, P.K., Li, J., and Lemaitre, B. (2026). Drosophila host defense mechanisms against filamentous fungal pathogens with diverse lifestyles. PLoS Pathog., 22.","DOI":"10.15252\/rc.2026268439"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2721","DOI":"10.1038\/s41467-026-68575-1","article-title":"The insect Toll pathway activates antibacterial immunity against the citrus Huanglongbing pathogen","volume":"17","author":"Du","year":"2026","journal-title":"Nat. Commun."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Jia, D., Luo, G., Guan, H., Yu, T., Sun, X., Du, Y., Wang, Y., Chen, H., and Wei, T. (2024). Arboviruses antagonize insect Toll antiviral immune signaling to facilitate the coexistence of viruses with their vectors. PLoS Pathog., 20.","DOI":"10.1371\/journal.ppat.1012318"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1111\/imb.12498","article-title":"Silencing of Apis mellifera dorsal genes reveals their role in expression of the antimicrobial peptide defensin-1","volume":"27","author":"Florecki","year":"2018","journal-title":"Insect Mol. Biol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Dong, Y., Taylor, H.E., and Dimopoulos, G. (2006). AgDscam, a hypervariable immunoglobulin domain-containing receptor of the Anopheles gambiae innate immune system. PLoS Biol., 4.","DOI":"10.1371\/journal.pbio.0040229"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1016\/j.chom.2012.09.004","article-title":"Anopheles NF-\u03baB-regulated splicing factors direct pathogen-specific repertoires of the hypervariable pattern recognition receptor AgDscam","volume":"12","author":"Dong","year":"2012","journal-title":"Cell Host Microbe"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Fan, Y., Yao, D., Ma, J., You, F., Wei, X., and Ji, T. (2024). Alternative splicing and alternative polyadenylation-regulated cold stress response of Apis cerana. Insects, 15.","DOI":"10.3390\/insects15121006"},{"key":"ref_25","first-page":"183","article-title":"Construction and annotation of the full-length transcriptome of the larval gut of Apis cerana cerana (Hymenoptera: Apidae) workers","volume":"67","author":"Song","year":"2024","journal-title":"Acta Entomol. Sin."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/j.coviro.2014.03.010","article-title":"The long and short of antiviral defense: Small RNA-based immunity in insects","volume":"7","author":"Bronkhorst","year":"2014","journal-title":"Curr. Opin. Virol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"6448","DOI":"10.1038\/s41598-017-06623-z","article-title":"Virus and dsRNA-triggered transcriptional responses reveal key components of honey bee antiviral defense","volume":"7","author":"Brutscher","year":"2017","journal-title":"Sci. Rep."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"880","DOI":"10.1111\/j.1462-5822.2006.00688.x","article-title":"RNAi is an antiviral immune response against a dsRNA virus in Drosophila melanogaster","volume":"8","author":"Zambon","year":"2006","journal-title":"Cell. Microbiol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"657","DOI":"10.1128\/MMBR.67.4.657-685.2003","article-title":"RNA interference: Biology, mechanism, and applications","volume":"67","author":"Agrawal","year":"2003","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"1699","DOI":"10.1002\/ps.5692","article-title":"Development of a Bacillus thuringiensis-based dsRNA production platform to control sacbrood virus in Apis cerana","volume":"76","author":"Park","year":"2020","journal-title":"Pest Manag. Sci."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1016\/S0092-8674(00)80620-0","article-title":"RNAi: Double-stranded RNA directs the ATP-dependent cleavage of mRNA at 21 to 23 nucleotide intervals","volume":"101","author":"Zamore","year":"2000","journal-title":"Cell"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"40","DOI":"10.1016\/j.gene.2017.04.022","article-title":"Uncovering the immune responses of Apis mellifera ligustica larval gut to Ascosphaera apis infection utilizing transcriptome sequencing","volume":"621","author":"Chen","year":"2017","journal-title":"Gene"},{"key":"ref_33","first-page":"208","article-title":"Identification and analysis of microRNAs in the larval gut of Apis cerana cerana","volume":"55","author":"Feng","year":"2022","journal-title":"Sci. Agric. Sin."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1038\/nbt.1754","article-title":"Integrative Genomics Viewer","volume":"29","author":"Robinson","year":"2011","journal-title":"Nat. Biotechnol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"509","DOI":"10.1515\/znc-2013-11-1210","article-title":"Molecular characterization and oxidative stress response of a cytochrome P450 gene (CYP4G11) from Apis cerana cerana","volume":"68","author":"Shi","year":"2013","journal-title":"Z. Naturforsch. C J. Biosci."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/j.biochi.2017.10.022","article-title":"Isolation of carboxylesterase (esterase FE4) from Apis cerana cerana and its role in oxidative resistance during adverse environmental stress","volume":"144","author":"Ma","year":"2018","journal-title":"Biochimie"},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis of relative gene expression data using real-time quantitative PCR and the 2\u2212\u0394\u0394CT method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"634","DOI":"10.1093\/clinchem\/hvaf043","article-title":"MIQE 2.0: Revision of the Minimum Information for Publication of Quantitative Real-Time PCR Experiments Guidelines","volume":"71","author":"Bustin","year":"2025","journal-title":"Clin. Chem."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"267","DOI":"10.5483\/BMBRep.2008.41.4.267","article-title":"Positive and negative regulation of the Drosophila immune response","volume":"41","author":"Aggarwal","year":"2008","journal-title":"BMB Rep."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"697","DOI":"10.1146\/annurev.immunol.25.022106.141615","article-title":"The host defense of Drosophila melanogaster","volume":"25","author":"Lemaitre","year":"2007","journal-title":"Annu. Rev. Immunol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"1869","DOI":"10.1242\/dev.128.10.1869","article-title":"Activation and repression by the C-terminal domain of Dorsal","volume":"128","author":"Jia","year":"2001","journal-title":"Development"}],"container-title":["Biomolecules"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2218-273X\/16\/9\/1223\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,25]],"date-time":"2026-08-25T04:31:53Z","timestamp":1787632313000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2218-273X\/16\/9\/1223"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,23]]},"references-count":41,"journal-issue":{"issue":"9","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["biom16091223"],"URL":"https:\/\/doi.org\/10.3390\/biom16091223","relation":{},"ISSN":["2218-273X"],"issn-type":[{"value":"2218-273X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,23]]}}}