{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,20]],"date-time":"2026-04-20T12:53:58Z","timestamp":1776689638319,"version":"3.51.2"},"reference-count":47,"publisher":"The Company of Biologists","issue":"6","license":[{"start":{"date-parts":[[2024,3,15]],"date-time":"2024-03-15T00:00:00Z","timestamp":1710460800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.biologists.com\/user-licence-1-1\/"}],"funder":[{"DOI":"10.13039\/100001654","name":"Grass Foundation","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100001654","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/100007185","name":"University of California, Los Angeles","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100007185","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2024,3,15]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n               <jats:p>Both male and female ticks have a strong innate drive to find and blood-feed on hosts. Carbon dioxide (CO2) is considered a critical behavioral activator and attractant for ticks and an essential sensory cue to find hosts. Yet, how CO2 activates and promotes host seeking in ticks is poorly understood. CO2 responses were studied in the black-legged tick Ixodes scapularis, the primary vector for Lyme disease in North America. Adult males and females were exposed to 1%, 2%, 4% or 8% CO2, and changes in walking behavior and foreleg movement were analyzed. CO2 is a potent stimulant for adult I. scapularis, even at lower concentrations (1%). Behavioral reactions depended on the animal's state: walking ticks increased their walking speed, while stationary ticks started to wave their forelegs and began to quest \u2013 both behaviors resembling aspects of host seeking. Only in sporadic cases did stationary animals start to walk when exposed to CO2, supporting the hypothesis that CO2 acts as an activator rather than an attractant. Furthermore, I. scapularis did not show a clear concentration preference and was not tuned more robustly to breath-like CO2 concentrations (\u223c4%) than to the other concentrations tested. Moreover, convincing evidence is provided showing that the foreleg Haller's organ is not necessary for CO2 detection. Even with a disabled or amputated Haller's organ, I. scapularis responded robustly to CO2, signifying that there must be CO2-sensitive structures important for tick host seeking that have not yet been identified.<\/jats:p>","DOI":"10.1242\/jeb.246874","type":"journal-article","created":{"date-parts":[[2024,2,19]],"date-time":"2024-02-19T09:44:45Z","timestamp":1708335885000},"source":"Crossref","is-referenced-by-count":7,"title":["The black-legged tick <i>Ixodes scapularis<\/i> detects CO2 without the Haller's organ"],"prefix":"10.1242","volume":"227","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1761-6296","authenticated-orcid":false,"given":"Carola","family":"St\u00e4dele","sequence":"first","affiliation":[{"name":"Institute for Neuro- and Sensory Physiology; University of G\u00f6ttingen Medical Center , G\u00f6ttingen, Lower Saxony 37073 , Germany"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"237","published-online":{"date-parts":[[2024,3,19]]},"reference":[{"key":"2024040907165785000_JEB246874C1","doi-asserted-by":"publisher","first-page":"649","DOI":"10.1016\/J.TINS.2016.08.002","article-title":"Smelling time: a neural basis for olfactory scene analysis","volume":"39","author":"Ache","year":"2016","journal-title":"Trends Neurosci."},{"key":"2024040907165785000_JEB246874C2","doi-asserted-by":"publisher","first-page":"63","DOI":"10.12688\/WELLCOMEOPENRES.15191.2","article-title":"Raincloud plots: a multi-platform tool for robust data visualization","volume":"4","author":"Allen","year":"2021","journal-title":"Wellcome Open Res."},{"key":"2024040907165785000_JEB246874C3","first-page":"59","article-title":"Life cycles and natural history of ticks","volume-title":"Biology of Ticks","author":"Apanaskevich","year":"2014"},{"key":"2024040907165785000_JEB246874C4","doi-asserted-by":"crossref","DOI":"10.1007\/978-3-662-04899-3","volume-title":"A Spider's World: Senses and Behavior","author":"Barth","year":"2002"},{"key":"2024040907165785000_JEB246874C5","doi-asserted-by":"publisher","first-page":"e0221659","DOI":"10.1371\/journal.pone.0221659","article-title":"Ticks home in on body heat: a new understanding of Haller's organ and repellent action","volume":"14","author":"Carr","year":"2019","journal-title":"PLoS One"},{"key":"2024040907165785000_JEB246874C6","doi-asserted-by":"publisher","first-page":"1563","DOI":"10.3390\/ijms18071563","article-title":"Tick Haller's organ, a new paradigm for arthropod olfaction: how ticks differ from insects","volume":"18","author":"Carr","year":"2017","journal-title":"Int. J. Mol. Sci."},{"key":"2024040907165785000_JEB246874C7","doi-asserted-by":"publisher","first-page":"427","DOI":"10.1016\/J.PT.2012.07.004","article-title":"Chemosensory behaviors of parasites","volume":"28","author":"Chaisson","year":"2012","journal-title":"Trends Parasitol."},{"key":"2024040907165785000_JEB246874C8","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/S10827-021-00798-1\/FIGURES\/1","article-title":"Active sensing in a dynamic olfactory world","volume":"50","author":"Crimaldi","year":"2022","journal-title":"J. Comput. Neurosci."},{"key":"2024040907165785000_JEB246874C9","doi-asserted-by":"publisher","first-page":"2963","DOI":"10.1242\/JEB.01736","article-title":"Carbon dioxide instantly sensitizes female yellow fever mosquitoes to human skin odours","volume":"208","author":"Dekker","year":"2005","journal-title":"J. Exp. Biol."},{"key":"2024040907165785000_JEB246874C10","doi-asserted-by":"publisher","first-page":"131","DOI":"10.1007\/S003590100183","article-title":"The antennal motor system of the stick insect Carausius morosus: anatomy and antennal movement pattern during walking","volume":"187","author":"D\u00fcrr","year":"2001","journal-title":"J. Comp. Physiol. A"},{"key":"2024040907165785000_JEB246874C11","doi-asserted-by":"publisher","first-page":"151","DOI":"10.1017\/S0007485300051221","article-title":"Host location by Aedes aegypti (Diptera: Culicidae): a wind tunnel study of chemical cues","volume":"81","author":"Eiras","year":"1991","journal-title":"Bull. Entomol. Res."},{"key":"2024040907165785000_JEB246874C12","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1016\/J.PT.2017.12.006","article-title":"The blacklegged tick, Ixodes scapularis: an increasing public health concern","volume":"34","author":"Eisen","year":"2018","journal-title":"Trends Parasitol."},{"key":"2024040907165785000_JEB246874C13","doi-asserted-by":"publisher","first-page":"22","DOI":"10.1007\/BF00339463\/METRICS","article-title":"Fine structure of tarsal sensilla in the tick Amblyomma americanum (L.)","volume":"114","author":"Foelix","year":"1971","journal-title":"Z. Zellforsch Mikrosk. Anat"},{"key":"2024040907165785000_JEB246874C14","doi-asserted-by":"publisher","first-page":"605","DOI":"10.1093\/AESA\/55.5.605","article-title":"Carbon dioxide as an attractant for certain ticks (Acarina: Argasidae and Ixodidae)","volume":"55","author":"Garcia","year":"1962","journal-title":"Ann. Entomol. Soc. Am."},{"key":"2024040907165785000_JEB246874C15","doi-asserted-by":"publisher","first-page":"1090","DOI":"10.4269\/ajtmh.1965.14.1090","article-title":"Collection of Dermacentor andersoni (Stiles) with carbon dioxide and its application in studies of Colorado tick fever virus","volume":"14","author":"Garcia","year":"1965","journal-title":"Am. J. Trop. Med. Hyg."},{"key":"2024040907165785000_JEB246874C16","doi-asserted-by":"publisher","first-page":"286","DOI":"10.1093\/JMEDENT\/6.3.286","article-title":"Reaction of the winter tick, Dermacentor albipictus (Packard), to CO2","volume":"6","author":"Garcia","year":"1969","journal-title":"J. Med. Entomol."},{"key":"2024040907165785000_JEB246874C17","doi-asserted-by":"publisher","first-page":"294","DOI":"10.3390\/INSECTS14030294","article-title":"Current knowledge on chemosensory-related candidate molecules potentially involved in tick olfaction via Haller's organ","volume":"14","author":"Gebremedhin","year":"2023","journal-title":"Insects"},{"key":"2024040907165785000_JEB246874C18","doi-asserted-by":"publisher","first-page":"617","DOI":"10.1093\/JMEDENT\/44.4.617","article-title":"Age-related changes in female mosquito carbon dioxide detection","volume":"44","author":"Grant","year":"2007","journal-title":"J. Med. Entomol."},{"key":"2024040907165785000_JEB246874C19","doi-asserted-by":"publisher","first-page":"35","DOI":"10.1007\/BF01262198","article-title":"A carbon dioxide trap for prolonged sampling of Ixodes ricinus L. populations","volume":"1","author":"Gray","year":"1985","journal-title":"Exp. Appl. Acarol."},{"key":"2024040907165785000_JEB246874C20","doi-asserted-by":"publisher","first-page":"2325","DOI":"10.1007\/S00436-012-3088-2\/TABLES\/1","article-title":"Comparative efficiency of two models of CO2 traps in the collection of free-living stages of Ixodides","volume":"111","author":"Guedes","year":"2012","journal-title":"Parasitol. Res."},{"key":"2024040907165785000_JEB246874C21","first-page":"123","article-title":"Attraction to carbon dioxide of unfed stages of Amblyomma triguttatum triguttatum Koch, under field conditions","volume":"26","author":"Guglielmone","year":"1985","journal-title":"Acarologia"},{"key":"2024040907165785000_JEB246874C22","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1016\/J.CUB.2011.01.048","article-title":"A sensory code for host seeking in parasitic nematodes","volume":"21","author":"Hallem","year":"2011","journal-title":"Curr. Biol."},{"key":"2024040907165785000_JEB246874C23","doi-asserted-by":"publisher","first-page":"331","DOI":"10.1111\/J.1365-2915.1995.TB00143.X","article-title":"Activation of Anopheles gambiae mosquitoes by carbon dioxide and human breath","volume":"9","author":"Healy","year":"1995","journal-title":"Med. Vet. Entomol."},{"key":"2024040907165785000_JEB246874C24","doi-asserted-by":"publisher","first-page":"195","DOI":"10.1007\/S003590050106\/METRICS","article-title":"Involvement of the suboesophageal and thoracic ganglia in the control of antennal movements in crickets","volume":"181","author":"Horseman","year":"1997","journal-title":"J. Comp. Physiol. A Sensory Neural Behav. Physiol."},{"key":"2024040907165785000_JEB246874C25","first-page":"439","article-title":"The ecology of Dermacentor albipidus (Packard)","volume":"4","author":"Howell","year":"1940","journal-title":"Proc. 6th Pacific Sci. Congr. 1939"},{"key":"2024040907165785000_JEB246874C26","doi-asserted-by":"publisher","first-page":"547","DOI":"10.1093\/JME\/TJX232","article-title":"Morphometric analysis of chemoreception organ in male and female ticks (Acari: Ixodidae)","volume":"55","author":"Josek","year":"2018","journal-title":"J. Med. Entomol."},{"key":"2024040907165785000_JEB246874C27","doi-asserted-by":"publisher","first-page":"104175","DOI":"10.1016\/J.JINSPHYS.2020.104175","article-title":"Neurophysiological and behavioral responses of blacklegged ticks to host odors","volume":"128","author":"Josek","year":"2021","journal-title":"J. Insect Physiol."},{"key":"2024040907165785000_JEB246874C28","doi-asserted-by":"publisher","first-page":"297","DOI":"10.1093\/JMEDENT\/33.3.297","article-title":"Ixodes (Ixodes) scapularis (Acari: Ixodidae): redescription of all active stages, distribution, hosts, geographical variation, and medical and veterinary importance","volume":"33","author":"Keirans","year":"1996","journal-title":"J. Med. Entomol."},{"key":"2024040907165785000_JEB246874C29","first-page":"340","article-title":"Electron microscopy studies of Haller's organ in the tick Ixodes persulcatus (Ixodidae)","volume":"11","author":"Leonovich","year":"1977","journal-title":"Parasitologiya"},{"key":"2024040907165785000_JEB246874C30","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1007\/BF01193968","article-title":"Regeneration of Haller's sensory organ in the tick Ixodes ricinus L","volume":"15","author":"Leonovich","year":"1992","journal-title":"Exp. Appl. Acarol."},{"key":"2024040907165785000_JEB246874C31","doi-asserted-by":"publisher","first-page":"2250","DOI":"10.1016\/J.CUB.2019.06.001","article-title":"General visual and contingent thermal cues interact to elicit attraction in female Aedes aegypti mosquitoes","volume":"29","author":"Liu","year":"2019","journal-title":"Curr. Biol."},{"key":"2024040907165785000_JEB246874C32","doi-asserted-by":"publisher","first-page":"689573","DOI":"10.3389\/FNBEH.2021.689573\/BIBTEX","article-title":"An inexpensive, high-precision, modular spherical treadmill setup optimized for Drosophila experiments","volume":"15","author":"Loesche","year":"2021","journal-title":"Front. Behav. Neurosci."},{"key":"2024040907165785000_JEB246874C33","doi-asserted-by":"publisher","first-page":"311","DOI":"10.1007\/S00114-002-0317-Z","article-title":"Attraction of the tropical bont tick, Amblyomma variegatum, to human breath and to the breath components acetone, NO and CO2","volume":"89","author":"McMahon","year":"2002","journal-title":"Naturwissenschaften"},{"key":"2024040907165785000_JEB246874C34","doi-asserted-by":"publisher","first-page":"1060","DOI":"10.1016\/j.cell.2013.12.044","article-title":"Multimodal integration of carbon dioxide and other sensory cues drives mosquito attraction to humans","volume":"156","author":"McMeniman","year":"2014","journal-title":"Cell"},{"key":"2024040907165785000_JEB246874C35","doi-asserted-by":"publisher","first-page":"491","DOI":"10.1093\/JMEDENT\/5.4.491","article-title":"A carbon dioxide bait trap for collecting ticks and fleas from animal burrows","volume":"5","author":"Miles","year":"1968","journal-title":"J. Med. Entomol."},{"key":"2024040907165785000_JEB246874C36","doi-asserted-by":"publisher","first-page":"764","DOI":"10.1016\/j.ttbdis.2017.06.001","article-title":"Infrared light detection by the Haller's organ of adult American dog ticks, Dermacentor variabilis (Ixodida: Ixodidae)","volume":"8","author":"Mitchell","year":"2017","journal-title":"Ticks Tick Borne Dis."},{"key":"2024040907165785000_JEB246874C37","first-page":"59","article-title":"The role of carbon dioxide as stimulant and attractant to the sand tampan, Ornithodoroa savignyi (Audouin)","volume":"31","author":"Nevill","year":"1964","journal-title":"Onderstepoort J. Vet. Res."},{"key":"2024040907165785000_JEB246874C38","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1007\/BF01270457","article-title":"Field sampling of unfed adults of Amblyomma hebraeum Koch","volume":"3","author":"Norval","year":"1987","journal-title":"Exp. Appl. Acarol."},{"key":"2024040907165785000_JEB246874C39","doi-asserted-by":"publisher","first-page":"295","DOI":"10.1016\/S0304-4017(00)00322-8","article-title":"Population dynamics of the free-living stages of Amblyomma cajennense (Fabricius, 1787) (Acari: Ixodidae) on pastures of Pedro Leopoldo, Minas Gerais State, Brazil","volume":"92","author":"Oliveira","year":"2000","journal-title":"Vet. Parasitol."},{"key":"2024040907165785000_JEB246874C40","doi-asserted-by":"publisher","first-page":"e72196","DOI":"10.7554\/ELIFE.72196","article-title":"Learning to predict target location with turbulent odor plumes","volume":"11","author":"Rigolli","year":"2022","journal-title":"Elife"},{"key":"2024040907165785000_JEB246874C41","volume-title":"Biology of Ticks","author":"Soneshine","year":"2014"},{"key":"2024040907165785000_JEB246874C42","doi-asserted-by":"publisher","first-page":"665","DOI":"10.1007\/BF00198976","article-title":"Perception of breath components by the tropical bont tick, Amblyomma variegatum Fabricius (Ixodidae). I. CO2-excited and CO2-inhibited receptors","volume":"170","author":"Steullet","year":"1992","journal-title":"J. Comp. Physiol. A"},{"key":"2024040907165785000_JEB246874C43","doi-asserted-by":"publisher","first-page":"220","DOI":"10.1111\/MVE.12214","article-title":"Behavioural responses of Ixodes ricinus nymphs to carbon dioxide and rodent odour","volume":"31","author":"van Duijvendijk","year":"2017","journal-title":"Med. Vet. Entomol."},{"key":"2024040907165785000_JEB246874C44","doi-asserted-by":"publisher","first-page":"2509","DOI":"10.1016\/J.CUB.2019.06.043","article-title":"Visual-olfactory integration in the human disease vector mosquito Aedes aegypti","volume":"29","author":"Vinauger","year":"2019","journal-title":"Curr. Biol."},{"key":"2024040907165785000_JEB246874C45","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1111\/J.1440-6055.1977.TB00064.X","article-title":"The sensory nervous system of the adult cattle tick Boophilus microplus (Canestrini) Ixodidae Part II. Scanning electron microscopy","volume":"16","author":"Waladde","year":"1977","journal-title":"Aust. J. Entomol."},{"key":"2024040907165785000_JEB246874C46","doi-asserted-by":"publisher","first-page":"301","DOI":"10.1016\/J.CUB.2012.12.038","article-title":"Drosophila tracks carbon dioxide in flight","volume":"23","author":"Wasserman","year":"2013","journal-title":"Curr. Biol."},{"key":"2024040907165785000_JEB246874C47","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1093\/JMEDENT\/9.3.245","article-title":"Chemo-attraction in the lone star tick (Acarina: Ixodidae). I. Response of different developmental stages to carbon dioxide administered via traps","volume":"9","author":"Wilson","year":"1972","journal-title":"J. Med. Entomol."}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.246874\/3368064\/jeb246874.pdf","content-type":"application\/pdf","content-version":"am","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.246874\/3415185\/jeb246874.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/journals.biologists.com\/jeb\/article-pdf\/doi\/10.1242\/jeb.246874\/3415185\/jeb246874.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,4,9]],"date-time":"2024-04-09T07:17:09Z","timestamp":1712647029000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/227\/6\/jeb246874\/345075\/The-black-legged-tick-Ixodes-scapularis-detects"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2024,3,15]]},"references-count":47,"journal-issue":{"issue":"6","published-print":{"date-parts":[[2024,3,15]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.246874","relation":{},"ISSN":["0022-0949","1477-9145"],"issn-type":[{"value":"0022-0949","type":"print"},{"value":"1477-9145","type":"electronic"}],"subject":[],"published":{"date-parts":[[2024,3,15]]},"article-number":"jeb246874"}}