{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,16]],"date-time":"2026-06-16T20:54:39Z","timestamp":1781643279078,"version":"3.54.5"},"reference-count":76,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,3,16]],"date-time":"2026-03-16T00:00:00Z","timestamp":1773619200000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,4,27]],"date-time":"2026-04-27T00:00:00Z","timestamp":1777248000000},"content-version":"vor","delay-in-days":42,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/100000060","name":"National Institute of Allergy and Infectious Diseases","doi-asserted-by":"publisher","award":["Z01 AI000810"],"award-info":[{"award-number":["Z01 AI000810"]}],"id":[{"id":"10.13039\/100000060","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000060","name":"National Institute of Allergy and Infectious Diseases","doi-asserted-by":"publisher","award":["ZIAAI001337"],"award-info":[{"award-number":["ZIAAI001337"]}],"id":[{"id":"10.13039\/100000060","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["BMC Genomics"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:sec>\n                    <jats:title>Background<\/jats:title>\n                    <jats:p>\n                      The lone star tick,\n                      <jats:italic>Amblyomma americanum<\/jats:italic>\n                      , is a three-host species widely distributed across North America and a vector for multiple human pathogens. Bites from this tick is also associated with alpha-gal syndrome, underscoring its growing public health importance. Adult female ticks undergo a prolonged feeding cycle divided into preparatory, slow-feeding, and rapid-feeding phases. Tick salivary glands are essential for successful blood feeding, producing a complex mixture of molecules that modulate host hemostasis, immunity, and tissue repair. The dynamic remodeling of the salivary gland transcriptome and proteome, termed the sialome switch, is thought to support prolonged feeding and immune evasion. To investigate the molecular mechanisms underlying the sialome switch, we performed high-throughput transcriptomic profiling of salivary glands from\n                      <jats:italic>A. americanum<\/jats:italic>\n                      adult females across the feeding cycle.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      Unlike previous studies that described the sialome switch in\n                      <jats:italic>A. americanum<\/jats:italic>\n                      by pooling ticks according to time post-attachment, we classified individuals based on average body weight, defining seven feeding stages encompassing unfed, slow-feeding, and rapid-feeding ticks. We identified 18,704 putative coding sequences (CDS) and revealed five distinct transcriptional profiles corresponding to different feeding stages. Unsupervised clustering indicated that most CDS were stage-specifically expressed, with temporal shifts observed in different protein families, including Kunitz-type, lipocalins, metalloproteases, and evasins. The transition from unfed to the slow-feeding phase corresponded to expression of a distinct subset of CDS within the signal transduction class, suggesting potential signaling pathways responsible for regulating this dynamic process.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Conclusions<\/jats:title>\n                    <jats:p>\n                      This study provides a high-resolution view of transcriptional dynamics in\n                      <jats:italic>A. americanum<\/jats:italic>\n                      salivary glands and identifies candidate molecular pathways regulating the sialome switch. These findings advance understanding of tick feeding biology and shed light on strategies ticks use to evade host defenses during prolonged blood meals.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.1186\/s12864-026-12666-2","type":"journal-article","created":{"date-parts":[[2026,3,16]],"date-time":"2026-03-16T12:33:04Z","timestamp":1773664384000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Further insights into the sialome switch of Amblyomma americanum adult females"],"prefix":"10.1186","volume":"27","author":[{"given":"Stephen","family":"Lu","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jose M.C.","family":"Ribeiro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lucas","family":"Tirloni","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,3,16]]},"reference":[{"issue":"2","key":"12666_CR1","doi-asserted-by":"publisher","first-page":"342","DOI":"10.1603\/ME13115","volume":"51","author":"YP Springer","year":"2014","unstructured":"Springer YP, Eisen L, Beati L, James AM, Eisen RJ. Spatial distribution of counties in the continental United States with records of occurrence of Amblyomma americanum (Ixodida: Ixodidae). J Med Entomol. 2014;51(2):342\u201351.","journal-title":"J Med Entomol"},{"key":"12666_CR2","doi-asserted-by":"publisher","unstructured":"SonenshineDE. Range Expansion of Tick Disease Vectors in North America: Implications for Spread of Tick-Borne Disease. Int J Environ Res Public Health. 2018;15(3). https:\/\/doi.org\/10.3390\/ijerph15030478.","DOI":"10.3390\/ijerph15030478"},{"issue":"2","key":"12666_CR3","first-page":"72","volume":"26","author":"CE Hopla","year":"1953","unstructured":"Hopla CE, Downs CM. The isolation of Bacterium tularense from the tick, Amblyomma americanum. J Kansas Entomol Soc. 1953;26(2):72\u20133.","journal-title":"J Kansas Entomol Soc"},{"issue":"12","key":"12666_CR4","doi-asserted-by":"publisher","first-page":"1810","DOI":"10.1086\/320721","volume":"183","author":"AM James","year":"2001","unstructured":"James AM, Liveris D, Wormser GP, Schwartz I, Montecalvo MA, Johnson BJ. Borrelia lonestari infection after a bite by an Amblyomma americanum tick. J Infect Dis. 2001;183(12):1810\u20134.","journal-title":"J Infect Dis"},{"issue":"2","key":"12666_CR5","doi-asserted-by":"publisher","first-page":"239","DOI":"10.4269\/ajtmh.1993.49.239","volume":"49","author":"BE Anderson","year":"1993","unstructured":"Anderson BE, Sims KG, Olson JG, Childs JE, Piesman JF, Happ CM, Maupin GO, Johnson BJ. Amblyomma americanum: a potential vector of human ehrlichiosis. Am J Trop Med Hyg. 1993;49(2):239\u201344.","journal-title":"Am J Trop Med Hyg"},{"issue":"6","key":"12666_CR6","doi-asserted-by":"publisher","first-page":"1183","DOI":"10.1093\/jme\/tjad060","volume":"60","author":"AP Dupuis 2nd","year":"2023","unstructured":"Dupuis AP 2nd, Lange RE, Ciota AT. Emerging tickborne viruses vectored by Amblyomma americanum (Ixodida: Ixodidae): Heartland and Bourbon viruses. J Med Entomol. 2023;60(6):1183\u201396.","journal-title":"J Med Entomol"},{"key":"12666_CR7","doi-asserted-by":"publisher","first-page":"775371","DOI":"10.3389\/fcimb.2021.775371","volume":"11","author":"SR Sharma","year":"2021","unstructured":"Sharma SR, Crispell G, Mohamed A, Cox C, Lange J, Choudhary S, Commins SP, Karim S. Alpha-Gal Syndrome: Involvement of Amblyomma americanum alpha-D-Galactosidase and beta-1,4 Galactosyltransferase Enzymes in alpha-Gal Metabolism. Front Cell Infect Microbiol. 2021;11:775371.","journal-title":"Front Cell Infect Microbiol"},{"key":"12666_CR8","unstructured":"Soneshine DE, Roe RM. Biology of Ticks. 2nd ed. Oxford University Press; 2013."},{"issue":"2","key":"12666_CR9","doi-asserted-by":"publisher","first-page":"332","DOI":"10.1084\/jem.161.2.332","volume":"161","author":"JM Ribeiro","year":"1985","unstructured":"Ribeiro JM, Makoul GT, Levine J, Robinson DR, Spielman A. Antihemostatic, antiinflammatory, and immunosuppressive properties of the saliva of a tick, Ixodes dammini. J Exp Med. 1985;161(2):332\u201344.","journal-title":"J Exp Med"},{"issue":"6","key":"12666_CR10","doi-asserted-by":"publisher","first-page":"2051","DOI":"10.2741\/3363","volume":"14","author":"IM Francischetti","year":"2009","unstructured":"Francischetti IM, Sa-Nunes A, Mans BJ, Santos IM, Ribeiro JM. The role of saliva in tick feeding. Front Biosci (Landmark Ed). 2009;14(6):2051\u201388.","journal-title":"Front Biosci (Landmark Ed)"},{"issue":"2","key":"12666_CR11","doi-asserted-by":"publisher","first-page":"97","DOI":"10.1016\/0020-7519(78)90004-8","volume":"8","author":"KC Binnington","year":"1978","unstructured":"Binnington KC. Sequential changes in salivary gland structure during attachment and feeding of the cattle tick, Boophilus microplus. Int J Parasitol. 1978;8(2):97\u2013115.","journal-title":"Int J Parasitol"},{"issue":"2","key":"12666_CR12","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/0020-7322(79)90007-2","volume":"8","author":"MJW Megaw","year":"1979","unstructured":"Megaw MJW, Beadle DJ. Structure and Function of the Salivary-Glands of the Tick, Boophilus-Microplus Canestrini (Acarina, Ixodidae). Int J Insect Morphol. 1979;8(2):67\u201383.","journal-title":"Int J Insect Morphol"},{"issue":"1","key":"12666_CR13","doi-asserted-by":"publisher","first-page":"100","DOI":"10.2307\/3281330","volume":"68","author":"JL McSwain","year":"1982","unstructured":"McSwain JL, Essenberg RC, Sauer JR. Protein changes in the salivary glands of the female lone star tick, Amblyomma americanum, during feeding. J Parasitol. 1982;68(1):100\u20136.","journal-title":"J Parasitol"},{"issue":"2","key":"12666_CR14","doi-asserted-by":"publisher","first-page":"e0007758","DOI":"10.1371\/journal.pntd.0007758","volume":"14","author":"TK Kim","year":"2020","unstructured":"Kim TK, Tirloni L, Pinto AFM, Diedrich JK, Moresco JJ, Yates JR 3rd, Vaz daS, Mulenga I Jr. Time-resolved proteomic profile of Amblyomma americanum tick saliva during feeding. PLoS Negl Trop Dis. 2020;14(2):e0007758.","journal-title":"PLoS Negl Trop Dis"},{"key":"12666_CR15","doi-asserted-by":"publisher","first-page":"374","DOI":"10.3389\/fcimb.2020.00374","volume":"10","author":"JMC Ribeiro","year":"2020","unstructured":"Ribeiro JMC, Mans BJ. TickSialoFam (TSFam): A Database That Helps to Classify Tick Salivary Proteins, a Review on Tick Salivary Protein Function and Evolution, With Considerations on the Tick Sialome Switching Phenomenon. Front Cell Infect Microbiol. 2020;10:374.","journal-title":"Front Cell Infect Microbiol"},{"issue":"2","key":"12666_CR16","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1016\/j.ttbdis.2018.12.001","volume":"10","author":"S Narasimhan","year":"2019","unstructured":"Narasimhan S, Booth CJ, DePonte K, Wu MJ, Liang X, Mohanty S, Kantor F, Fikrig E. Host-specific expression of Ixodes scapularis salivary genes. Ticks Tick Borne Dis. 2019;10(2):386\u201397.","journal-title":"Ticks Tick Borne Dis"},{"key":"12666_CR17","doi-asserted-by":"publisher","first-page":"517","DOI":"10.3389\/fcimb.2017.00517","volume":"7","author":"L Tirloni","year":"2017","unstructured":"Tirloni L, Kim TK, Pinto AFM, Yates JR 3rd, da Silva Vaz I Jr., Mulenga A. Tick-Host Range Adaptation: Changes in Protein Profiles in Unfed Adult Ixodes scapularis and Amblyomma americanum Saliva Stimulated to Feed on Different Hosts. Front Cell Infect Microbiol. 2017;7:517.","journal-title":"Front Cell Infect Microbiol"},{"issue":"2","key":"12666_CR18","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1111\/j.1365-2583.2009.00863.x","volume":"18","author":"MN Aljamali","year":"2009","unstructured":"Aljamali MN, Hern L, Kupfer D, Downard S, So S, Roe BA, Sauer JR, Essenberg RC. Transcriptome analysis of the salivary glands of the female tick Amblyomma americanum (Acari: Ixodidae). Insect Mol Biol. 2009;18(2):129\u201354.","journal-title":"Insect Mol Biol"},{"key":"12666_CR19","doi-asserted-by":"publisher","unstructured":"KarimS, Ribeiro JMC. An Insight into the Sialome of the Lone Star Tick, with a Glimpse on Its Time Dependent Gene Expression. PLoS ONE. 2015;10(7). https:\/\/doi.org\/10.1371\/journal.pone.0131292.","DOI":"10.1371\/journal.pone.0131292"},{"key":"12666_CR20","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1146\/annurev.ento.48.060402.102812","volume":"48","author":"JM Ribeiro","year":"2003","unstructured":"Ribeiro JM, Francischetti IM. Role of arthropod saliva in blood feeding: sialome and post-sialome perspectives. Annu Rev Entomol. 2003;48:73\u201388.","journal-title":"Annu Rev Entomol"},{"key":"12666_CR21","doi-asserted-by":"publisher","first-page":"518","DOI":"10.1186\/1471-2164-15-518","volume":"15","author":"ZM Radulovic","year":"2014","unstructured":"Radulovic ZM, Kim TK, Porter LM, Sze SH, Lewis L, Mulenga A. A 24\u201348\u00a0h fed Amblyomma americanum tick saliva immuno-proteome. BMC Genomics. 2014;15:518.","journal-title":"BMC Genomics"},{"key":"12666_CR22","unstructured":"Balashov YS. Bloodsucking Ticks (Ixodoidea)\u2014Vectors of Diseases of Man and Animals. 1st. ed. Nauka Publishers; 1968."},{"issue":"5","key":"12666_CR23","doi-asserted-by":"publisher","first-page":"732","DOI":"10.1093\/jmedent\/44.5.732","volume":"44","author":"DR Troughton","year":"2007","unstructured":"Troughton DR, Levin ML. Life cycles of seven ixodid tick species (Acari: Ixodidae) under standardized laboratory conditions. J Med Entomol. 2007;44(5):732\u201340.","journal-title":"J Med Entomol"},{"issue":"7","key":"12666_CR24","doi-asserted-by":"publisher","first-page":"644","DOI":"10.1038\/nbt.1883","volume":"29","author":"MG Grabherr","year":"2011","unstructured":"Grabherr MG, Haas BJ, Yassour M, Levin JZ, Thompson DA, Amit I, Adiconis X, Fan L, Raychowdhury R, Zeng Q, et al. Full-length transcriptome assembly from RNA-Seq data without a reference genome. Nat Biotechnol. 2011;29(7):644\u201352.","journal-title":"Nat Biotechnol"},{"issue":"6","key":"12666_CR25","doi-asserted-by":"publisher","first-page":"1117","DOI":"10.1101\/gr.089532.108","volume":"19","author":"JT Simpson","year":"2009","unstructured":"Simpson JT, Wong K, Jackman SD, Schein JE, Jones SJ. Birol I: ABySS: a parallel assembler for short read sequence data. Genome Res. 2009;19(6):1117\u201323.","journal-title":"Genome Res"},{"issue":"23","key":"12666_CR26","doi-asserted-by":"publisher","first-page":"3150","DOI":"10.1093\/bioinformatics\/bts565","volume":"28","author":"L Fu","year":"2012","unstructured":"Fu L, Niu B, Zhu Z, Wu S, Li W. CD-HIT: accelerated for clustering the next-generation sequencing data. Bioinformatics. 2012;28(23):3150\u20132.","journal-title":"Bioinformatics"},{"issue":"4","key":"12666_CR27","doi-asserted-by":"publisher","first-page":"783","DOI":"10.1016\/j.jmb.2004.05.028","volume":"340","author":"JD Bendtsen","year":"2004","unstructured":"Bendtsen JD, Nielsen H, von Heijne G, Brunak S. Improved prediction of signal peptides: SignalP 3.0. J Mol Biol. 2004;340(4):783\u201395.","journal-title":"J Mol Biol"},{"key":"12666_CR28","doi-asserted-by":"publisher","unstructured":"LiB, Dewey CN. RSEM: accurate transcript quantification from RNA-Seq data with or without a reference genome. BMC Bioinformatics 2011;12. https:\/\/doi.org\/10.1186\/1471-2105-12-323.","DOI":"10.1186\/1471-2105-12-323"},{"issue":"Database issue","key":"12666_CR29","doi-asserted-by":"publisher","first-page":"D222","DOI":"10.1093\/nar\/gku1221","volume":"43","author":"A Marchler-Bauer","year":"2015","unstructured":"Marchler-Bauer A, Derbyshire MK, Gonzales NR, Lu S, Chitsaz F, Geer LY, Geer RC, He J, Gwadz M, Hurwitz DI, et al. CDD: NCBI\u2019s conserved domain database. Nucleic Acids Res. 2015;43(Database issue):D222\u2013226.","journal-title":"Nucleic Acids Res"},{"key":"12666_CR30","doi-asserted-by":"publisher","unstructured":"KarimS, Singh P, Ribeiro JMC. A Deep Insight into the Sialotranscriptome of the Gulf Coast Tick, Amblyomma maculatum. PLoS ONE. 2011;6(12). https:\/\/doi.org\/10.1371\/journal.pone.0028525.","DOI":"10.1371\/journal.pone.0028525"},{"issue":"19","key":"12666_CR31","doi-asserted-by":"publisher","first-page":"3210","DOI":"10.1093\/bioinformatics\/btv351","volume":"31","author":"FA Simao","year":"2015","unstructured":"Simao FA, Waterhouse RM, Ioannidis P, Kriventseva EV, Zdobnov EM. BUSCO: assessing genome assembly and annotation completeness with single-copy orthologs. Bioinformatics. 2015;31(19):3210\u20132.","journal-title":"Bioinformatics"},{"issue":"1","key":"12666_CR32","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1093\/bioinformatics\/btp616","volume":"26","author":"MD Robinson","year":"2010","unstructured":"Robinson MD, McCarthy DJ, Smyth GK. edgeR: a Bioconductor package for differential expression analysis of digital gene expression data. Bioinformatics. 2010;26(1):139\u201340.","journal-title":"Bioinformatics"},{"key":"12666_CR33","unstructured":"R Core Team. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria; 2025. https:\/\/www.R-project.org\/."},{"key":"12666_CR34","doi-asserted-by":"publisher","first-page":"232","DOI":"10.1186\/1471-2105-6-232","volume":"6","author":"R Shamir","year":"2005","unstructured":"Shamir R, Maron-Katz A, Tanay A, Linhart C, Steinfeld I, Sharan R, Shiloh Y, Elkon R. EXPANDER\u2013an integrative program suite for microarray data analysis. BMC Bioinformatics. 2005;6:232.","journal-title":"BMC Bioinformatics"},{"issue":"13","key":"12666_CR35","doi-asserted-by":"publisher","first-page":"1710","DOI":"10.1093\/bioinformatics\/btg213","volume":"19","author":"DP Wall","year":"2003","unstructured":"Wall DP, Fraser HB, Hirsh AE. Detecting putative orthologs. Bioinformatics. 2003;19(13):1710\u20131.","journal-title":"Bioinformatics"},{"key":"12666_CR36","doi-asserted-by":"publisher","first-page":"103899","DOI":"10.1016\/j.jprot.2020.103899","volume":"229","author":"L Tirloni","year":"2020","unstructured":"Tirloni L, Lu S, Calvo E, Sabadin G, Di Maggio LS, Suzuki M, Nardone G, da Silva Vaz I Jr., Ribeiro JMC. Integrated analysis of sialotranscriptome and sialoproteome of the brown dog tick Rhipicephalus sanguineus (s.l.): Insights into gene expression during blood feeding. J Proteom. 2020;229:103899.","journal-title":"J Proteom"},{"issue":"1","key":"12666_CR37","doi-asserted-by":"publisher","first-page":"996","DOI":"10.1186\/s12864-024-10905-y","volume":"25","author":"S Lu","year":"2024","unstructured":"Lu S, de Sousa-Paula LC, Ribeiro JMC, Tirloni L. Exploring the longitudinal expression dynamics of midguts in adult female Amblyomma americanum ticks. BMC Genomics. 2024;25(1):996.","journal-title":"BMC Genomics"},{"issue":"11","key":"12666_CR38","doi-asserted-by":"publisher","first-page":"1261","DOI":"10.1016\/S0022-1910(01)00112-3","volume":"47","author":"BL Weiss","year":"2001","unstructured":"Weiss BL, Reuben Kaufman W. The relationship between \u2018critical weight\u2019 and 20-hydroxyecdysone in the female ixodid tick, Amblyomma hebraeum. J Insect Physiol. 2001;47(11):1261\u20137.","journal-title":"J Insect Physiol"},{"issue":"D1","key":"12666_CR39","doi-asserted-by":"publisher","first-page":"D624","DOI":"10.1093\/nar\/gkx1134","volume":"46","author":"ND Rawlings","year":"2018","unstructured":"Rawlings ND, Barrett AJ, Thomas PD, Huang X, Bateman A, Finn RD. The MEROPS database of proteolytic enzymes, their substrates and inhibitors in 2017 and a comparison with peptidases in the PANTHER database. Nucleic Acids Res. 2018;46(D1):D624\u201332.","journal-title":"Nucleic Acids Res"},{"issue":"36","key":"12666_CR40","doi-asserted-by":"publisher","first-page":"26298","DOI":"10.1074\/jbc.M513010200","volume":"281","author":"M Kotsyfakis","year":"2006","unstructured":"Kotsyfakis M, Sa-Nunes A, Francischetti IM, Mather TN, Andersen JF, Ribeiro JM. Antiinflammatory and immunosuppressive activity of sialostatin L, a salivary cystatin from the tick Ixodes scapularis. J Biol Chem. 2006;281(36):26298\u2013307.","journal-title":"J Biol Chem"},{"key":"12666_CR41","doi-asserted-by":"publisher","first-page":"288","DOI":"10.3389\/fcimb.2017.00288","volume":"7","author":"T Zavasnik-Bergant","year":"2017","unstructured":"Zavasnik-Bergant T, Vidmar R, Sekirnik A, Fonovic M, Salat J, Grunclova L, Kopacek P, Turk B. Salivary Tick Cystatin OmC2 Targets Lysosomal Cathepsins S and C in Human Dendritic Cells. Front Cell Infect Microbiol. 2017;7:288.","journal-title":"Front Cell Infect Microbiol"},{"issue":"10","key":"12666_CR42","doi-asserted-by":"publisher","first-page":"3602","DOI":"10.1182\/blood-2001-12-0237","volume":"99","author":"IM Francischetti","year":"2002","unstructured":"Francischetti IM, Valenzuela JG, Andersen JF, Mather TN, Ribeiro JM. Ixolaris, a novel recombinant tissue factor pathway inhibitor (TFPI) from the salivary gland of the tick, Ixodes scapularis: identification of factor X and factor Xa as scaffolds for the inhibition of factor VIIa\/tissue factor complex. Blood. 2002;99(10):3602\u201312.","journal-title":"Blood"},{"issue":"4","key":"12666_CR43","doi-asserted-by":"publisher","first-page":"318","DOI":"10.1016\/j.exppara.2009.04.007","volume":"122","author":"C Stutzer","year":"2009","unstructured":"Stutzer C, Mans BJ, Gaspar AR, Neitz AW, Maritz-Olivier C. Ornithodoros savignyi: soft tick apyrase belongs to the 5\u2019-nucleotidase family. Exp Parasitol. 2009;122(4):318\u201327.","journal-title":"Exp Parasitol"},{"issue":"3-4","key":"12666_CR44","doi-asserted-by":"publisher","first-page":"211","DOI":"10.1016\/j.ijpara.2017.08.018","volume":"48","author":"T Hollmann 3rd","year":"2018","unstructured":"Hollmann T, Kim TK, Tirloni L, Radulovic ZM, Pinto AFM, Diedrich JK, Yates JR 3rd, da, Silva Vaz I, Mulenga A Jr. Identification and characterization of proteins in the Amblyomma americanum tick cement cone. Int J Parasitol. 2018, 48(3\u20134):211\u2013224.","journal-title":"Int J Parasitol"},{"issue":"1","key":"12666_CR45","doi-asserted-by":"publisher","first-page":"21300","DOI":"10.1038\/s41598-022-24881-4","volume":"12","author":"A Mulenga","year":"2022","unstructured":"Mulenga A, Radulovic Z, Porter L, Britten TH, Kim TK, Tirloni L, Gaithuma AK, Adeniyi-Ipadeola GO, Dietrich JK, Moresco JJ, et al. Identification and characterization of proteins that form the inner core Ixodes scapularis tick attachment cement layer. Sci Rep. 2022;12(1):21300.","journal-title":"Sci Rep"},{"issue":"4","key":"12666_CR46","doi-asserted-by":"publisher","first-page":"e0006410","DOI":"10.1371\/journal.pntd.0006410","volume":"12","author":"J Perner","year":"2018","unstructured":"Perner J, Kropackova S, Kopacek P, Ribeiro JMC. Sialome diversity of ticks revealed by RNAseq of single tick salivary glands. PLoS Negl Trop Dis. 2018;12(4):e0006410.","journal-title":"PLoS Negl Trop Dis"},{"issue":"3","key":"12666_CR47","doi-asserted-by":"publisher","first-page":"727","DOI":"10.1242\/jeb.64.3.727","volume":"64","author":"W Kaufman","year":"1976","unstructured":"Kaufman W. The influence of various factors on fluid secretion by in vitro salivary glands of ixodid Ticks. J Exp Biol. 1976;64(3):727\u201342.","journal-title":"J Exp Biol"},{"issue":"1","key":"12666_CR48","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/0306-4492(82)90197-6","volume":"72","author":"SP Schmidt","year":"1982","unstructured":"Schmidt SP, Essenberg RC, Sauer JR. A dopamine sensitive adenylate cyclase in the salivary glands of Amblyomma americanum (L). Comp Biochem Physiol C Comp Pharmacol. 1982;72(1):9\u201314.","journal-title":"Comp Biochem Physiol C Comp Pharmacol"},{"issue":"1","key":"12666_CR49","doi-asserted-by":"publisher","first-page":"16054","DOI":"10.1038\/s41598-020-73077-1","volume":"10","author":"L Mateos-Hernandez","year":"2020","unstructured":"Mateos-Hernandez L, Defaye B, Vancova M, Hajdusek O, Sima R, Park Y, Attoui H, Simo L. Cholinergic axons regulate type I acini in salivary glands of Ixodes ricinus and Ixodes scapularis ticks. Sci Rep. 2020;10(1):16054.","journal-title":"Sci Rep"},{"issue":"10","key":"12666_CR50","doi-asserted-by":"publisher","first-page":"e76128","DOI":"10.1371\/journal.pone.0076128","volume":"8","author":"H Gomes","year":"2013","unstructured":"Gomes H, Romeiro NC, Braz GR, de Oliveira EA, Rodrigues C, da Fonseca RN, Githaka N, Isezaki M, Konnai S, Ohashi K, et al. Identification and structural-functional analysis of cyclin-dependent kinases of the cattle tick Rhipicephalus (Boophilus) microplus. PLoS ONE. 2013;8(10):e76128.","journal-title":"PLoS ONE"},{"issue":"1","key":"12666_CR51","doi-asserted-by":"publisher","first-page":"43","DOI":"10.1016\/S0965-1748(98)00103-9","volume":"29","author":"MJ Palmer","year":"1999","unstructured":"Palmer MJ, McSwain JL, Spatz MD, Tucker JS, Essenberg RC, Sauer JR. Molecular cloning of cAMP-dependent protein kinase catalytic subunit isoforms from the lone star tick, Amblyomma americanum (L). Insect Biochem Mol Biol. 1999;29(1):43\u201351.","journal-title":"Insect Biochem Mol Biol"},{"issue":"3","key":"12666_CR52","doi-asserted-by":"publisher","first-page":"340","DOI":"10.1006\/gcen.1995.1118","volume":"99","author":"H Mao","year":"1995","unstructured":"Mao H, McBlain WA, Kaufman WR. Some properties of the ecdysteroid receptor in the salivary gland of the ixodid tick, Amblyomma hebraeum. Gen Comp Endocrinol. 1995;99(3):340\u20138.","journal-title":"Gen Comp Endocrinol"},{"issue":"1","key":"12666_CR53","doi-asserted-by":"publisher","first-page":"386","DOI":"10.1186\/s13071-021-04879-z","volume":"14","author":"S Hu","year":"2021","unstructured":"Hu S, Wang Y, Xu Z, Zhou Y, Cao J, Zhang H, Zhou J. Identification of the Bcl-2 and Bax homologs from Rhipicephalus haemaphysaloides and their function in the degeneration of tick salivary glands. Parasit Vectors. 2021;14(1):386.","journal-title":"Parasit Vectors"},{"issue":"1","key":"12666_CR54","doi-asserted-by":"publisher","first-page":"612","DOI":"10.1186\/s13071-021-05052-2","volume":"14","author":"X Lu","year":"2021","unstructured":"Lu X, Zhang Z, Yuan D, Zhou Y, Cao J, Zhang H, da Silva Vaz I Jr., Zhou J. The ecdysteroid receptor regulates salivary gland degeneration through apoptosis in Rhipicephalus haemaphysaloides. Parasit Vectors. 2021;14(1):612.","journal-title":"Parasit Vectors"},{"issue":"4","key":"12666_CR55","doi-asserted-by":"publisher","first-page":"303","DOI":"10.1080\/15592294.2016.1163460","volume":"11","author":"A Cabezas-Cruz","year":"2016","unstructured":"Cabezas-Cruz A, Alberdi P, Ayllon N, Valdes JJ, Pierce R, Villar M, de la Fuente J. Anaplasma phagocytophilum increases the levels of histone modifying enzymes to inhibit cell apoptosis and facilitate pathogen infection in the tick vector Ixodes scapularis. Epigenetics. 2016;11(4):303\u201319.","journal-title":"Epigenetics"},{"issue":"6","key":"12666_CR56","doi-asserted-by":"publisher","first-page":"e0324546","DOI":"10.1371\/journal.pone.0324546","volume":"20","author":"GH MacIntosh","year":"2025","unstructured":"MacIntosh GH, Nuyens AC, Vickery JL, Berthold A, Lloyd VK. Epigenetic responses in Borrelia-infected Ixodes scapularis ticks: Over-expression of euchromatic histone lysine methyltransferase 2 and no change in DNA methylation. PLoS ONE. 2025;20(6):e0324546.","journal-title":"PLoS ONE"},{"key":"12666_CR57","doi-asserted-by":"publisher","unstructured":"AmaranteAM, de Oliveira DM, de Souza M, Fonseca-Oliveira M, Galina A, Rosignoli S, Arcanjo AF, Moraes B, Paiardini A, Rotili D et al. Unraveling the Roles of Epigenetic Regulators During the Embryonic Development of Rhipicephalus microplus. Int J Mol Sci. 2025;26(18). https:\/\/doi.org\/10.3390\/ijms26189171.","DOI":"10.3390\/ijms26189171"},{"key":"12666_CR58","doi-asserted-by":"publisher","first-page":"109933","DOI":"10.1016\/j.vetpar.2023.109933","volume":"318","author":"L Liu","year":"2023","unstructured":"Liu L, Cheng R, Mao SQ, Duan DY, Feng LL, Cheng TY. Saliva proteome of partially- and fully-engorged adult female Haemaphysalis flava ticks. Vet Parasitol. 2023;318:109933.","journal-title":"Vet Parasitol"},{"issue":"7","key":"12666_CR59","doi-asserted-by":"publisher","first-page":"927","DOI":"10.1017\/S0031182017001998","volume":"145","author":"BF Leal","year":"2018","unstructured":"Leal BF, Alzugaray MF, Seixas A, Da Silva Vaz I, Ferreira CAS. Characterization of a glycine-rich protein from Rhipicephalus microplus: tissue expression, gene silencing and immune recognition. Parasitology. 2018;145(7):927\u201338.","journal-title":"Parasitology"},{"issue":"5","key":"12666_CR60","doi-asserted-by":"publisher","first-page":"578","DOI":"10.1093\/jmedent\/32.5.578","volume":"32","author":"AN Alekseev","year":"1995","unstructured":"Alekseev AN, Burenkova LA, Podboronov VM, Chunikhin SP. Bacteriocidal qualities of ixodid tick (Acarina: Ixodidae) salivary cement plugs and their changes under the influence of a viral tick-borne pathogen. J Med Entomol. 1995;32(5):578\u201382.","journal-title":"J Med Entomol"},{"issue":"4","key":"12666_CR61","doi-asserted-by":"publisher","first-page":"1652","DOI":"10.1128\/IAI.67.4.1652-1658.1999","volume":"67","author":"A Mulenga","year":"1999","unstructured":"Mulenga A, Sugimoto C, Sako Y, Ohashi K, Musoke A, Shubash M, Onuma M. Molecular characterization of a Haemaphysalis longicornis tick salivary gland-associated 29-kilodalton protein and its effect as a vaccine against tick infestation in rabbits. Infect Immun. 1999;67(4):1652\u20138.","journal-title":"Infect Immun"},{"issue":"12","key":"12666_CR62","doi-asserted-by":"publisher","first-page":"1289","DOI":"10.1292\/jvms.68.1289","volume":"68","author":"T Harnnoi","year":"2006","unstructured":"Harnnoi T, Watchabunsook S, Sakaguchi T, Xuan X, Fujisaki K. Characterization of Haemaphysalis longicornis recombinant cement-like antigens and preliminary study of their vaccination effects. J Vet Med Sci. 2006;68(12):1289\u201395.","journal-title":"J Vet Med Sci"},{"issue":"1","key":"12666_CR63","doi-asserted-by":"publisher","first-page":"77","DOI":"10.1007\/s00436-006-0243-7","volume":"100","author":"J Zhou","year":"2006","unstructured":"Zhou J, Gong H, Zhou Y, Xuan X, Fujisaki K. Identification of a glycine-rich protein from the tick Rhipicephalus haemaphysaloides and evaluation of its vaccine potential against tick feeding. Parasitol Res. 2006;100(1):77\u201384.","journal-title":"Parasitol Res"},{"issue":"12","key":"12666_CR64","doi-asserted-by":"publisher","first-page":"7422","DOI":"10.4049\/jimmunol.0900075","volume":"182","author":"A Sa-Nunes","year":"2009","unstructured":"Sa-Nunes A, Bafica A, Antonelli LR, Choi EY, Francischetti IM, Andersen JF, Shi GP, Chavakis T, Ribeiro JM, Kotsyfakis M. The immunomodulatory action of sialostatin L on dendritic cells reveals its potential to interfere with autoimmunity. J Immunol. 2009;182(12):7422\u20139.","journal-title":"J Immunol"},{"issue":"10","key":"12666_CR65","doi-asserted-by":"publisher","first-page":"2003","DOI":"10.1007\/s00018-019-03034-3","volume":"76","author":"J Kotal","year":"2019","unstructured":"Kotal J, Stergiou N, Busa M, Chlastakova A, Berankova Z, Rezacova P, Langhansova H, Schwarz A, Calvo E, Kopecky J, et al. The structure and function of Iristatin, a novel immunosuppressive tick salivary cystatin. Cell Mol Life Sci. 2019;76(10):2003\u201313.","journal-title":"Cell Mol Life Sci"},{"issue":"5","key":"12666_CR66","doi-asserted-by":"publisher","first-page":"661","DOI":"10.1016\/S1097-2765(00)80359-7","volume":"3","author":"GC Paesen","year":"1999","unstructured":"Paesen GC, Adams PL, Harlos K, Nuttall PA, Stuart DI. Tick histamine-binding proteins: isolation, cloning, and three-dimensional structure. Mol Cell. 1999;3(5):661\u201371.","journal-title":"Mol Cell"},{"issue":"27","key":"12666_CR67","doi-asserted-by":"publisher","first-page":"18721","DOI":"10.1074\/jbc.M800188200","volume":"283","author":"BJ Mans","year":"2008","unstructured":"Mans BJ, Ribeiro JM, Andersen JF. Structure, function, and evolution of biogenic amine-binding proteins in soft ticks. J Biol Chem. 2008;283(27):18721\u201333.","journal-title":"J Biol Chem"},{"issue":"3","key":"12666_CR68","doi-asserted-by":"publisher","first-page":"784","DOI":"10.1016\/j.jmb.2007.03.064","volume":"369","author":"P Roversi","year":"2007","unstructured":"Roversi P, Lissina O, Johnson S, Ahmat N, Paesen GC, Ploss K, Boland W, Nunn MA, Lea SM. The structure of OMCI, a novel lipocalin inhibitor of the complement system. J Mol Biol. 2007;369(3):784\u201393.","journal-title":"J Mol Biol"},{"issue":"6","key":"12666_CR69","doi-asserted-by":"publisher","first-page":"102209","DOI":"10.1016\/j.ttbdis.2023.102209","volume":"14","author":"MG Guizzo","year":"2023","unstructured":"Guizzo MG, Mans B, Pienaar R, Ribeiro JMC. A comparison of Illumina and PacBio methods to build tick salivary gland transcriptomes confirms large expression of lipocalins and other salivary protein families that are not represented in available tick genomes. Ticks Tick Borne Dis. 2023;14(6):102209.","journal-title":"Ticks Tick Borne Dis"},{"issue":"4","key":"12666_CR70","doi-asserted-by":"publisher","first-page":"869","DOI":"10.1016\/S0006-291X(03)00857-X","volume":"305","author":"IM Francischetti","year":"2003","unstructured":"Francischetti IM, Mather TN, Ribeiro JM. Cloning of a salivary gland metalloprotease and characterization of gelatinase and fibrin(ogen)lytic activities in the saliva of the Lyme disease tick vector Ixodes scapularis. Biochem Biophys Res Commun. 2003;305(4):869\u201375.","journal-title":"Biochem Biophys Res Commun"},{"issue":"7","key":"12666_CR71","doi-asserted-by":"publisher","first-page":"1485","DOI":"10.1111\/j.1742-4658.2008.06308.x","volume":"275","author":"Y Decrem","year":"2008","unstructured":"Decrem Y, Beaufays J, Blasioli V, Lahaye K, Brossard M, Vanhamme L, Godfroid E. A family of putative metalloproteases in the salivary glands of the tick Ixodes ricinus. FEBS J. 2008;275(7):1485\u201399.","journal-title":"FEBS J"},{"key":"12666_CR72","doi-asserted-by":"publisher","first-page":"563349","DOI":"10.3389\/fcimb.2020.563349","volume":"10","author":"J Perner","year":"2020","unstructured":"Perner J, Helm D, Haberkant P, Hatalova T, Kropackova S, Ribeiro JM, Kopacek P. The Central Role of Salivary Metalloproteases in Host Acquired Resistance to Tick Feeding. Front Cell Infect Microbiol. 2020;10:563349.","journal-title":"Front Cell Infect Microbiol"},{"key":"12666_CR73","doi-asserted-by":"publisher","first-page":"769542","DOI":"10.3389\/fcimb.2021.769542","volume":"11","author":"S Bhattacharya","year":"2021","unstructured":"Bhattacharya S, Nuttall PA. Phylogenetic Analysis Indicates That Evasin-Like Proteins of Ixodid Ticks Fall Into Three Distinct Classes. Front Cell Infect Microbiol. 2021;11:769542.","journal-title":"Front Cell Infect Microbiol"},{"issue":"37","key":"12666_CR74","doi-asserted-by":"publisher","first-page":"27250","DOI":"10.1074\/jbc.M704706200","volume":"282","author":"A Frauenschuh","year":"2007","unstructured":"Frauenschuh A, Power CA, Deruaz M, Ferreira BR, Silva JS, Teixeira MM, Dias JM, Martin T, Wells TNC, Proudfoot AEI. Molecular cloning and characterization of a highly selective chemokine-binding protein from the tick Rhipicephalus sanguineus. J Biol Chem. 2007;282(37):27250\u20138.","journal-title":"J Biol Chem"},{"issue":"12","key":"12666_CR75","doi-asserted-by":"publisher","first-page":"e8514","DOI":"10.1371\/journal.pone.0008514","volume":"4","author":"JM Dias","year":"2009","unstructured":"Dias JM, Losberger C, Deruaz M, Power CA, Proudfoot AE, Shaw JP. Structural basis of chemokine sequestration by a tick chemokine binding protein: the crystal structure of the complex between Evasin-1 and CCL3. PLoS ONE. 2009;4(12):e8514.","journal-title":"PLoS ONE"},{"issue":"9","key":"12666_CR76","doi-asserted-by":"publisher","first-page":"2019","DOI":"10.1084\/jem.20072689","volume":"205","author":"M Deruaz","year":"2008","unstructured":"Deruaz M, Frauenschuh A, Alessandri AL, Dias JM, Coelho FM, Russo RC, Ferreira BR, Graham GJ, Shaw JP, Wells TN, et al. Ticks produce highly selective chemokine binding proteins with antiinflammatory activity. J Exp Med. 2008;205(9):2019\u201331.","journal-title":"J Exp Med"}],"container-title":["BMC Genomics"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/link.springer.com\/article\/10.1186\/s12864-026-12666-2","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12864-026-12666-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/link.springer.com\/content\/pdf\/10.1186\/s12864-026-12666-2.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,6,16]],"date-time":"2026-06-16T20:11:17Z","timestamp":1781640677000},"score":1,"resource":{"primary":{"URL":"https:\/\/link.springer.com\/10.1186\/s12864-026-12666-2"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,3,16]]},"references-count":76,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["12666"],"URL":"https:\/\/doi.org\/10.1186\/s12864-026-12666-2","relation":{},"ISSN":["1471-2164"],"issn-type":[{"value":"1471-2164","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,3,16]]},"assertion":[{"value":"12 December 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"13 February 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"16 March 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Animal experiments were conducted in accordance with the guidelines of the National Institutes of Health on protocols approved by the Rocky Mountain Laboratories Animal Care and Use Committee (2020-065 and 2023-037). The Rocky Mountain Veterinary Branch is accredited by the International Association for Assessment and Accreditation of Laboratory Animal Care (AAALAC).","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Ethics approval and consent to participate"}},{"value":"Not applicable.","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Consent for publication"}},{"value":"The authors declare no competing interests.","order":4,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}],"article-number":"411"}}