{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,1,23]],"date-time":"2026-01-23T12:31:02Z","timestamp":1769171462721,"version":"3.49.0"},"reference-count":57,"publisher":"Public Library of Science (PLoS)","issue":"1","license":[{"start":{"date-parts":[[2025,1,10]],"date-time":"2025-01-10T00:00:00Z","timestamp":1736467200000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100012705","name":"Szkola Gl\u00f3wna Gospodarstwa Wiejskiego w Warszawie","doi-asserted-by":"publisher","award":["The APC is financed by Warsaw University of Life Sciences. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript."],"award-info":[{"award-number":["The APC is financed by Warsaw University of Life Sciences. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript."]}],"id":[{"id":"10.13039\/501100012705","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["www.plosone.org"],"crossmark-restriction":false},"short-container-title":["PLoS ONE"],"abstract":"<jats:p>The study involved a gross anatomical description of the parotid gland, mandibular gland, monostomatic sublingual gland, polystomatic sublingual gland, and zygomatic gland in 12 adult Eurasian wolves (<jats:italic>Canis lupus lupus<\/jats:italic>) (wild free-ranging individuals and their zoo counterparts), including their morphometry and microscopic evaluation using hematoxylin &amp; eosin, mucicarmine, azan trichrome, PAS, AB pH 1.0, AB pH 2.5; AB pH 2.5\/PAS, and HDI stainings. Topographically, the salivary glands were located similarly to those of other terrestrial Carnivora. The parotid gland in all wolves had a shape similar to a trapezoid (four angles). The parotid duct opened onto the fourth upper premolar (P4). The parotid gland was a branched alveolar complex that produced serous secretion. In captive specimens, the mandibular gland was a complex branched tubular gland producing mucous secretion, while in free-ranging wolves it was a branched tubuloalveolar gland producing mucoserous secretion. The monostomatic sublingual gland in free-ranging wolves was a complex branched tubuloalveolar gland that produced seromucous secretion, while in captive wolves, it revealed a mucoserous secretion character. The polystomatic sublingual gland consisted of several independent packets (from 6\u20137 to 7\u20138) and was a complex branched tubuloalveolar gland with seromucous secretion. The zygomatic duct opened onto the last upper molar tooth (M3), and this gland was a complex branched tubular gland producing mucous secretion. The anatomical and histological similarities between the salivary glands of the oral cavity in captive and free-ranging wolves, compared to other terrestrial carnivores, provide valuable insights for veterinary treatments and understanding pathological conditions. These findings emphasize the need for further research on diverse populations of wolves and related species within the Canidae family to better understand the influence of diet on salivary gland morphology.<\/jats:p>","DOI":"10.1371\/journal.pone.0317102","type":"journal-article","created":{"date-parts":[[2025,1,10]],"date-time":"2025-01-10T20:04:51Z","timestamp":1736539491000},"page":"e0317102","update-policy":"https:\/\/doi.org\/10.1371\/journal.pone.corrections_policy","source":"Crossref","is-referenced-by-count":1,"title":["Anatomical variations in Eurasian wolf (Canis lupus lupus) (Carnivora: Canidae) of the salivary glands: A histological and histochemical investigation"],"prefix":"10.1371","volume":"20","author":[{"given":"Joanna","family":"Kle\u0107kowska-Nawrot","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4412-1966","authenticated-orcid":true,"given":"Karolina","family":"Go\u017adziewska-Har\u0142ajczuk","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Krzysztof","family":"Stegmann","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Arkadiusz","family":"Dziech","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Dariusz","family":"\u0141upicki","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Gabriela","family":"J\u0119drszczyk","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Igor","family":"Jucenco","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Karolina","family":"Barszcz","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2025,1,10]]},"reference":[{"key":"pone.0317102.ref001","volume-title":"Nomina Anatomica Veterinaria","year":"2017","edition":"6"},{"key":"pone.0317102.ref002","volume-title":"Textbook of Veterinary Anatomy.","author":"KM Dyce","year":"1996","edition":"2"},{"key":"pone.0317102.ref003","volume-title":"Lehrbuch der Anatomie der Haustiere","author":"R Nickel","year":"2004"},{"key":"pone.0317102.ref004","volume-title":"Glandular mechanisms of salivary secretion. Front Oral Biology","author":"B Tandler","year":"1998"},{"key":"pone.0317102.ref005","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1002\/ar.1108","article-title":"Secretion by striated ducts of mammalian major salivary glands: Review from an ultrastructural, functional, and evolutionary perspective","volume":"264","author":"B Tandler","year":"2001","journal-title":"Anat Rec"},{"issue":"4","key":"pone.0317102.ref006","first-page":"221","article-title":"Investigations on the shape and size of molar and zygomatic salivary glands in shorthair domestic cats","volume":"12","author":"AA Mohammadpour","year":"2009","journal-title":"Bulgarian J Vet Med"},{"key":"pone.0317102.ref007","doi-asserted-by":"crossref","first-page":"34","DOI":"10.3906\/vet-1709-36","article-title":"First evidence of histological and histochemical intraspecific differences in salivary glands in Brown Norway and albino Wistar rats","volume":"42","author":"F Ruxanda","year":"2018","journal-title":"Turkish J Vet Anim Sci"},{"issue":"2","key":"pone.0317102.ref008","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1067\/mpr.2001.113778","article-title":"A review of saliva: Normal composition, flow and function","volume":"85","author":"SP Humphrey","year":"2001","journal-title":"J Prosthet Dent"},{"key":"pone.0317102.ref009","doi-asserted-by":"crossref","first-page":"156","DOI":"10.1016\/j.tvjl.2015.07.017","article-title":"Computed tomography and magnetic resonance diagnosis of variations in the anatomical location of the major salivary glands in 1680 dogs and 187 cats","volume":"209","author":"A Durand","year":"2016","journal-title":"Vet J"},{"issue":"2","key":"pone.0317102.ref010","first-page":"252","article-title":"Surgical anatomy, morphometry, and histochemistry of major salivary glands in dogs: Updates and Recommendations","volume":"8","author":"W Gaber","year":"2020","journal-title":"Internat J Vet Health Sci Res"},{"key":"pone.0317102.ref011","doi-asserted-by":"crossref","first-page":"551","DOI":"10.1111\/ahe.12396","article-title":"Anatomic and magnetic resonance imaging features of the salivary glands in the dog","volume":"47","author":"F Gil","year":"2018","journal-title":"Anat Histol Embryol"},{"issue":"11","key":"pone.0317102.ref012","doi-asserted-by":"crossref","first-page":"637","DOI":"10.1369\/0022155417732527","article-title":"Canine salivary glands: Analysis of Rab and SNARE protein expression and SNARE complex formation with diverse tissue properties","volume":"65","author":"H Gomi","year":"2017","journal-title":"J Histochem Cytochem"},{"key":"pone.0317102.ref013","volume-title":"Miller and Evans\u2019 Anatomy of the Dog","author":"JW Hermanson","year":"2020","edition":"5"},{"key":"pone.0317102.ref014","author":"LCU Junqueira","year":"1965"},{"key":"pone.0317102.ref015","doi-asserted-by":"crossref","first-page":"397","DOI":"10.1111\/j.1439-0264.2011.01084.x","article-title":"CT sialography in the dog\u2013A cadaver study","volume":"40","author":"S Kneissl","year":"2011","journal-title":"Anat Histol Embryol"},{"issue":"1","key":"pone.0317102.ref016","doi-asserted-by":"crossref","first-page":"1","DOI":"10.21608\/jva.2022.226444","article-title":"Anatomical-Surgical study on the thyro-laryngeal region of Native dogs (Canis lupus familiaris)","volume":"15","author":"MA Nazih","year":"2022","journal-title":"J Vet Anat"},{"key":"pone.0317102.ref017","doi-asserted-by":"crossref","first-page":"269","DOI":"10.1007\/BF01839193","article-title":"Localization of glycoconjugates in dog parotid gland by lectin histochemistry","volume":"18","author":"V Pedini","year":"1994","journal-title":"Vet Res Comm"},{"issue":"3","key":"pone.0317102.ref018","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1016\/0034-5288(94)90130-9","article-title":"Glycoconjugates in the mandibular salivary gland of adult dogs revealed by lectin histochemistry","volume":"57","author":"V Pedini","year":"1994","journal-title":"Res Vet Sci"},{"key":"pone.0317102.ref019","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1007\/BF01839318","article-title":"A lectin histochemical study of the zygomatic salivary gland of adult dogs","volume":"19","author":"V Pedini","year":"1995","journal-title":"Vet Res Comm"},{"key":"pone.0317102.ref020","first-page":"285","article-title":"IgA, IgG and IgM immunohistochemical staining in normal dog salivary glands and in cases of chronic sialadenitis","volume":"27","author":"M Sozmen","year":"1996","journal-title":"Vet Res"},{"key":"pone.0317102.ref021","first-page":"583","article-title":"Lectin histochemistry of dog major and minor salivary glands","volume":"30","author":"M Sozmen","year":"1999","journal-title":"Vet Res"},{"key":"pone.0317102.ref022","first-page":"99","article-title":"Quantitation of histochemical staining of salivary gland mucin using analysis in cats and dogs","volume":"30","author":"M Sozmen","year":"1999","journal-title":"Vet Res"},{"issue":"3","key":"pone.0317102.ref023","first-page":"225","article-title":"Sialography in dog: normal appearance","volume":"74","author":"M Tadjalli","year":"2004","journal-title":"Vet Archiv"},{"issue":"2","key":"pone.0317102.ref024","doi-asserted-by":"crossref","first-page":"149","DOI":"10.1111\/j.1439-0264.2011.01115.x","article-title":"MR anatomy of salivary glands in the dog","volume":"41","author":"S Weidner","year":"2012","journal-title":"Anat Histol Embryol"},{"issue":"7","key":"pone.0317102.ref025","first-page":"85","article-title":"Histological study of salivary glands in cats and dogs","volume":"3","author":"C Zhang","year":"2022","journal-title":"MEDS Clinic Medi"},{"issue":"4","key":"pone.0317102.ref026","first-page":"414","article-title":"Anatomical, Histological and Radiological Studies of Parotid Salivary Glands of the Red Fox (Vulpes vulpes) with Comparison to Baladi Dog","volume":"11","author":"AM Ibrahium","year":"2022","journal-title":"Internat J Vet Sci"},{"key":"pone.0317102.ref027","doi-asserted-by":"crossref","first-page":"e37528","DOI":"10.4025\/actascibiolsci.v40i1.37528","article-title":"Anatomical description of the salivary glands: parotid, mandibular and sublingual of the crab-eating-fox (Cerdocyon thous)","volume":"40","author":"TSB Pereira","year":"2018","journal-title":"Acta Scien. Biol Sci"},{"issue":"1\u20132","key":"pone.0317102.ref028","doi-asserted-by":"crossref","first-page":"23","DOI":"10.4322\/rbcv.2016.024","article-title":"Topography of the parotid duct opening in Cerdocyon thous (Linnaeus, 1766) and Lycalopex gymnocercus (G. Fischer, 1814)","volume":"23","author":"WV Souza","year":"2016","journal-title":"Revista Brasileira de Ci\u00eancia Vet"},{"issue":"2","key":"pone.0317102.ref029","first-page":"333","article-title":"Lima EMMD. Topography of the openings of the parotid ducts of a \u201cGuara\u201d wolf (Chrysocyon brachyurus Illiger, 1811)","volume":"31","author":"ALQ Santos","year":"2001","journal-title":"Ci\u00eancia Rural"},{"key":"pone.0317102.ref030","doi-asserted-by":"crossref","first-page":"457","DOI":"10.1186\/s12917-024-04305-3","article-title":"The comparative morphology of the oral cavity of glands in the captive South African painted dog (Lycaon pictus pictus) and fennec fox (Vulpes zerda) (Carnivora: Canide)","volume":"20","author":"J Kle\u0107kowska-Nawrot","year":"2024","journal-title":"BMC Vet Res"},{"issue":"6","key":"pone.0317102.ref031","doi-asserted-by":"crossref","first-page":"234","DOI":"10.1111\/j.1748-5827.1997.tb03353.x","article-title":"Lipomatous infiltration of the canine salivary gland","volume":"38","author":"J Brown P","year":"1997","journal-title":"J Small Anim Pract"},{"key":"pone.0317102.ref032","first-page":"647","volume-title":"Manual Saunders small animal clinic","author":"SE Johnson","year":"2008"},{"issue":"2","key":"pone.0317102.ref033","doi-asserted-by":"crossref","first-page":"173","DOI":"10.5455\/OVJ.2015.v5.i2.p173","article-title":"Ultrasound and multidetector computed tomography of mandibular salivary gland adenocarcinoma in two dogs","volume":"5","author":"D Lenoci","year":"2015","journal-title":"Open Vet J"},{"issue":"4","key":"pone.0317102.ref034","doi-asserted-by":"crossref","first-page":"604","DOI":"10.1177\/1040638720932169","article-title":"A retrospective study of salivary gland diseases in 179 dogs (2010\u20132018)","volume":"32","author":"DE Lieske","year":"2020","journal-title":"J Vet Diagn Invest"},{"issue":"3","key":"pone.0317102.ref035","doi-asserted-by":"crossref","first-page":"465","DOI":"10.2460\/javma.1991.198.03.465","article-title":"Salivary gland disease in dogs and cats: 245 cases (1985\u20131988)","volume":"198","author":"WL Spangler","year":"1991","journal-title":"J Am Vet Med Assoc"},{"issue":"2","key":"pone.0317102.ref036","doi-asserted-by":"crossref","first-page":"473","DOI":"10.7589\/2018-05-127","article-title":"Isolation of rabies virus from the salivary glands of wild and domestic carnivores during a skunk rabies epizootic","volume":"55","author":"I Jimenez","year":"2019","journal-title":"J Wildlife Diseas"},{"key":"pone.0317102.ref037","volume-title":"Angewandte und Topographische Anatomie der Haustiere","author":"R. Berg","year":"1973"},{"key":"pone.0317102.ref038","volume-title":"Technika histologiczna.","author":"H-C Burck","year":"1975"},{"key":"pone.0317102.ref039","first-page":"78","volume-title":"Series on Methods and Achievements in Experimental Pathology","author":"SS Spicer","year":"1967"},{"key":"pone.0317102.ref040","volume-title":"Nomina Histologica Veterinaria.","year":"2017"},{"key":"pone.0317102.ref041","volume-title":"Biblioteka Przyrodnicza i \u0141owiecka","author":"Okarma H. Wilk","year":"2015"},{"key":"pone.0317102.ref042","volume-title":"Wolves, ecology and conservation","author":"RO Peterson","year":"2003"},{"issue":"4","key":"pone.0317102.ref043","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1111\/mam.12067","article-title":"Food habits of the world\u2019s grey wolves","volume":"46","author":"TM Newsome","year":"2016","journal-title":"Mammal Rev"},{"issue":"4","key":"pone.0317102.ref044","doi-asserted-by":"crossref","first-page":"335","DOI":"10.4098\/AT.arch.95-35","article-title":"The tropic ecology of wolves and their predatory role in ungulate communities of forest ecosystems in Europe","volume":"40","author":"H. Okarma","year":"1995","journal-title":"Acta Theriol"},{"key":"pone.0317102.ref045","doi-asserted-by":"crossref","first-page":"27","DOI":"10.1007\/BF00317805","article-title":"Wolf predation and snow cover as mortality factors in the ungulate community of the Bia\u0142owie\u017ca National Park, Poland.","volume":"90","author":"W J\u0119drzejewski","year":"1992","journal-title":"Oecologia"},{"key":"pone.0317102.ref046","doi-asserted-by":"crossref","first-page":"709","DOI":"10.1016\/j.mambio.2011.06.007","article-title":"Diet and prey selection of wolves (Canis lupus) recolonising Western and Central Poland","volume":"76","author":"S Nowak","year":"2011","journal-title":"Mammal Biol"},{"issue":"1","key":"pone.0317102.ref047","doi-asserted-by":"crossref","first-page":"283","DOI":"10.2193\/2006-368","article-title":"Wolf depredation on domestic animals in the Polish Carpathian Mountains","volume":"72","author":"R. Gula","year":"2008","journal-title":"J Wildlife Managem"},{"issue":"2","key":"pone.0317102.ref048","doi-asserted-by":"crossref","first-page":"263","DOI":"10.1007\/BF03194489","article-title":"Patterns of wolf Canis lupus predation on wild and domestic ungulates in the Western Carpathian Mountains (S Poland)","volume":"50","author":"S Nowak","year":"2005","journal-title":"Acta Theriol"},{"key":"pone.0317102.ref049","first-page":"1","volume-title":"Analysis of the current types and extent of damage caused by wolves (Canis lupus) and the use of methods for resolving conflict situations","author":"S Nowak","year":"2005"},{"key":"pone.0317102.ref050","volume-title":"Large Canid (Canidae) Care Manual. Association of ZOO & Aquariums.","author":"M Rodden","year":"2012"},{"key":"pone.0317102.ref051","volume-title":"Anatomia zwierza Tom 2","author":"K Krysiak","year":"2012"},{"issue":"1","key":"pone.0317102.ref052","first-page":"81","article-title":"Anatomy and morphometry of major salivary glands of domestic cats with relation to their histological features","volume":"42","author":"YS Abouelela","year":"2022","journal-title":"Pakistan Vet J"},{"key":"pone.0317102.ref053","first-page":"174","article-title":"Aspetti ultrastrutturali di cellule sierose, mucose e \u2019siero-mucose","volume":"89","author":"G Aureli","year":"1966","journal-title":"La Clinica Vet"},{"key":"pone.0317102.ref054","volume-title":"chapter: 58: Sisson and Grossman\u2019s The anatomy of the domestic animals","author":"C. Diesem","year":"1975"},{"issue":"2","key":"pone.0317102.ref055","first-page":"223","article-title":"The labial and zygomatic salivary glands in mixed breed dogs in Trinidad: Anatomical location, histological features and histochemical characteristics","volume":"10","author":"R. Mohamed","year":"2020","journal-title":"World\u2019s Vet J"},{"key":"pone.0317102.ref056","first-page":"1","article-title":"Gross and microscopic characterization of the zygomatic salivary gland of the Philippine non-descript dog","volume":"45","author":"CP Maala","year":"2008","journal-title":"Philippine J Vet Med"},{"issue":"4","key":"pone.0317102.ref057","doi-asserted-by":"crossref","first-page":"730","DOI":"10.2307\/2261642","article-title":"Phylogeny and ecology reconsidered","volume":"83","author":"DD Ackerly","year":"1995","journal-title":"J Ecol"}],"container-title":["PLOS ONE"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0317102","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,1,10]],"date-time":"2025-01-10T20:05:26Z","timestamp":1736539526000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pone.0317102"}},"subtitle":[],"editor":[{"given":"Serkan","family":"Erdo\u011fan","sequence":"first","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2025,1,10]]},"references-count":57,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2025,1,10]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pone.0317102","relation":{},"ISSN":["1932-6203"],"issn-type":[{"value":"1932-6203","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,1,10]]}}}