{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T15:14:54Z","timestamp":1789139694954,"version":"build-2803163510"},"reference-count":127,"publisher":"MDPI AG","issue":"18","license":[{"start":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T00:00:00Z","timestamp":1789084800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>The domestic dog is the most morphologically diverse domesticated mammalian species. That diversity has arisen from deliberate, sustained artificial selection over the most recent 1% of the domestic dog\u2019s evolutionary history. However, artificial selection has pushed many individual breeds so far from the ancestral canine form that many dogs within these breeds can no longer perform the critical biological functions their bodies evolved to serve. Indeed, much of the popularity of many dog breeds is predicated on them no longer being functional dogs; rather, they are representations or caricatures of human aesthetic desires. The result is breeds in which the teleonome of individual dogs, i.e., the integrated biological system by which they maintain viability has been structurally dismantled in the pursuit of appearance. This article applies teleonomic and allostatic load theory to a multi-domain analysis of extreme brachycephaly as a primary case study, drawing on representative examples from other extreme phenotypes to examine their welfare costs across conformational disorder, innate health, allostatic load, and veterinary costs. A detailed Five Domains analysis of extreme brachycephalic morphotypes, where the epidemiological evidence is most complete, reveals that every major welfare domain is simultaneously compromised from birth because artificial selection has structurally compromised teleonomic function across morphological, physiological, physical, and behavioural traits. Allostatic load theory then provides the mechanistic explanation for the resulting shorter lifespans. The article concludes by addressing the veterinarian\u2019s emerging role as arbiter of animals\u2019 fitness for purpose under consumer and welfare laws, making the case for an Innate Health Assessment to become a standardised functional assessment framework grounded in teleonomic principles.<\/jats:p>","DOI":"10.3390\/ani16182867","type":"journal-article","created":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T14:39:24Z","timestamp":1789137564000},"page":"2867","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["\u2018Teleonomes in Tatters\u2019: The Welfare Costs of Selecting Dogs for Extreme Phenotypes, and the Case for Change"],"prefix":"10.3390","volume":"16","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7220-8378","authenticated-orcid":false,"given":"Paul D.","family":"McGreevy","sequence":"first","affiliation":[{"name":"Faculty of Science, Sydney School of Veterinary Science, University of Sydney, Sydney, NSW 2006, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"David J.","family":"Mellor","sequence":"additional","affiliation":[{"name":"Animal Welfare Science and Bioethics Centre, School of Veterinary Science, Massey University, Palmerston North 4442, New Zealand"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1115-2723","authenticated-orcid":false,"given":"Dan","family":"O\u2019Neill","sequence":"additional","affiliation":[{"name":"Veterinary Epidemiology, Economics and Public Health, The Royal Veterinary College, Hawkshead Lane, North Mymms, Hatfield AL9 7TA, Herts, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-9988-9828","authenticated-orcid":false,"given":"Rowena M.","family":"Packer","sequence":"additional","affiliation":[{"name":"Department of Clinical Science and Services, The Royal Veterinary College, Hawkshead Lane, North Mymms, Hatfield AL9 7TA, Herts, UK"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cathrynne","family":"Henshall","sequence":"additional","affiliation":[{"name":"School Agricultural, Environmental and Veterinary Sciences, Southern Cross University, Lismore, NSW 2678, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-4222-1727","authenticated-orcid":false,"given":"Cristina L.","family":"Wilkins","sequence":"additional","affiliation":[{"name":"School of Environmental and Rural Sciences, University of New England, Armidale, NSW 2351, Australia"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,11]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"105662","DOI":"10.1016\/j.biosystems.2025.105662","article-title":"On the origin of biological teleonomy","volume":"259","author":"Deng","year":"2026","journal-title":"BioSystems"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"341","DOI":"10.1093\/biolinnean\/blad037","article-title":"Teleonomy in living systems: An overview","volume":"139","author":"Corning","year":"2023","journal-title":"Biol. J. Linn. Soc."},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Wilkins, C.L., Henshall, C., Lykins, A.D., Mellor, D.J., Fillios, M., and McGreevy, P.D. (2026). The teleonome: A framework for understanding animal welfare integrating adaptive capabilities, affective regulation, agency, and environmental affordances. Front. Anim. Sci., 7.","DOI":"10.3389\/fanim.2026.1768519"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"606","DOI":"10.1016\/j.tvjl.2013.09.009","article-title":"How does severe brachycephaly affect dogs\u2019 lives? Results of a structured preoperative owner questionnaire","volume":"198","author":"Roedler","year":"2013","journal-title":"Vet. J."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Packer, R.M.A., Hendricks, A., Tivers, M.S., and Burn, C.C. (2015). Impact of facial conformation on canine health: Brachycephalic obstructive airway syndrome. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0137496"},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Packer, R.M.A., Hendricks, A., and Burn, C.C. (2015). Impact of facial conformation on canine health: Corneal ulceration. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0123827"},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Packer, R.M.A., O\u2019Neill, D.G., Fletcher, F., and Farnworth, M.J. (2019). Great expectations, inconvenient truths, and the paradoxes of the dog-owner relationship for owners of brachycephalic dogs. PLoS ONE, 14.","DOI":"10.1371\/journal.pone.0219918"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"17251","DOI":"10.1038\/s41598-020-73088-y","article-title":"Unravelling the health status of brachycephalic dogs in the UK using multivariable analysis","volume":"10","author":"Pegram","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1186\/s40575-021-00112-3","article-title":"French Bulldogs differ to other dogs in the UK in propensity for many common disorders: A VetCompass study","volume":"8","author":"Packer","year":"2021","journal-title":"Canine Med. Genet."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1186\/s40575-022-00118-5","article-title":"English Bulldogs in the UK: A VetCompass study of their disorder predispositions and protections","volume":"9","author":"Skipper","year":"2022","journal-title":"Canine Med. Genet."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"4","DOI":"10.1186\/s40575-022-00117-6","article-title":"Health of Pug dogs in the UK: Disorder predispositions and protections","volume":"9","author":"Sahota","year":"2022","journal-title":"Canine Med. Genet."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"1135","DOI":"10.1111\/jvim.17019","article-title":"Evaluation of risk factors for sleep-disordered breathing in dogs","volume":"38","author":"Niinikoski","year":"2024","journal-title":"J. Vet. Intern. Med."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Wilkins, C.L., Henshall, C., Lykins, A., Mellor, D.J., Fillios, M., and McGreevy, P.D. (2026). Applying the Teleonome: A biologically grounded process for welfare assessment that operationalises animal agency. Front. Anim. Sci., 7.","DOI":"10.3389\/fanim.2026.1768519"},{"key":"ref_14","unstructured":"Shaw, R., and Bransford, J. (1977). The Theory of Affordances. Perceiving, Acting, and Knowing: Toward an Ecological Psychology, Lawrence Erlbaum."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1097","DOI":"10.3758\/s13423-016-1183-6","article-title":"Doggone affordances: Canine perception of affordances for reaching","volume":"24","author":"Wagman","year":"2017","journal-title":"Psychon. Bull. Rev."},{"key":"ref_16","doi-asserted-by":"crossref","unstructured":"Seeley, K.E., Proudfoot, K.L., and Edes, A.N. (2022). The application of allostasis and allostatic load in animal species: A scoping review. PLoS ONE, 17.","DOI":"10.1371\/journal.pone.0273838"},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"104672","DOI":"10.1016\/j.neubiorev.2022.104672","article-title":"Mapping the brain basis of feelings, emotions and much more: A special issue focused on \u2018The Human Affectome\u2019","volume":"137","author":"Cromwell","year":"2022","journal-title":"Neurosci. Biobehav. Rev."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"705","DOI":"10.1038\/nrg.2017.67","article-title":"Demographic history, selection and functional diversity of the canine genome","volume":"18","author":"Ostrander","year":"2017","journal-title":"Nat. Rev. Genet."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1233","DOI":"10.1093\/shm\/hkz049","article-title":"The \u201cDog Doctors\u201d of Edwardian London: Elite canine veterinary care in the early twentieth century","volume":"33","author":"Skipper","year":"2020","journal-title":"Soc. Hist. Med."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"329","DOI":"10.1017\/S0962728600021965","article-title":"Some practical solutions to welfare problems in dog breeding","volume":"8","author":"McGreevy","year":"1999","journal-title":"Anim. Welf."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"125","DOI":"10.1017\/S0962728600031821","article-title":"Breeding for quality of life","volume":"16","author":"McGreevy","year":"2007","journal-title":"Anim. Welf."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1186\/2052-6687-1-2","article-title":"Approaches to canine health surveillance","volume":"1","author":"Church","year":"2014","journal-title":"Canine Genet. Epidemiol."},{"key":"ref_23","unstructured":"(2026, January 31). VetCompass. VetCompass Programme. Available online: https:\/\/www.rvc.ac.uk\/VetCOMPASS\/."},{"key":"ref_24","unstructured":"(2026, May 17). VetCompass Australia. VetCompass Australia. Available online: http:\/\/www.vetcompass.com.au\/."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"1300","DOI":"10.2460\/javma.240.11.1300","article-title":"Incidence of intervertebral disk degeneration-related diseases and associated mortality rates in dogs","volume":"240","author":"Bergknut","year":"2012","journal-title":"J. Am. Vet. Med. Assoc."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Bannasch, D., Batcher, K., Leuthard, F., Bannasch, M., Hug, P., Marcellin-Little, D.J., Dickinson, P.J., Dr\u00f6gem\u00fcller, M., Dr\u00f6gem\u00fcller, C., and Leeb, T. (2022). The effects of FGF4 retrogenes on canine morphology. Genes, 13.","DOI":"10.3390\/genes13020325"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"e17281","DOI":"10.1111\/jvim.17281","article-title":"The relationship between radiographic disc calcification score and FGF4L2 genotype in dachshunds","volume":"39","author":"Sullivan","year":"2025","journal-title":"J. Vet. Intern. Med."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1016\/j.tvjl.2009.08.033","article-title":"Inherited defects in pedigree dogs. Part 1: Disorders related to breed standards","volume":"182","author":"Asher","year":"2009","journal-title":"Vet. J."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"638","DOI":"10.1016\/j.tvjl.2013.09.020","article-title":"Longevity and mortality of owned dogs in England","volume":"198","author":"Church","year":"2013","journal-title":"Vet. J."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1111\/jsap.12723","article-title":"Gastric dilation-volvulus in dogs attending UK emergency-care veterinary practices: Prevalence, risk factors and survival","volume":"58","author":"Case","year":"2017","journal-title":"J. Small Anim. Pract."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1186\/s40575-023-00131-2","article-title":"Dog breeds and conformations predisposed to osteosarcoma in the UK: A VetCompass study","volume":"10","author":"Edmunds","year":"2023","journal-title":"Canine Med. Genet."},{"key":"ref_32","first-page":"99","article-title":"Chiari-like malformation in the Cavalier King Charles Spaniel","volume":"42","author":"Rusbridge","year":"2006","journal-title":"J. Am. Anim. Hosp. Assoc."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1136\/vr.103651","article-title":"Dogs attending primary-care practice in England with clinical signs suggestive of Chiari-like malformation\/syringomyelia","volume":"179","author":"Pelligand","year":"2016","journal-title":"Vet. Rec."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Brambilla, P.G., Polli, M., Pradelli, D., Papa, M., Rizzi, R., Bagardi, M., and Bussadori, C. (2020). Epidemiological study of congenital heart diseases in dogs: Prevalence, popularity, and volatility throughout twenty years of clinical practice. PLoS ONE, 15.","DOI":"10.1101\/2020.02.25.964262"},{"key":"ref_35","unstructured":"Maggs, D.J., Miller, P.E., and Ofri, R. (2018). Diseases of the eyelids. Slatter\u2019s Fundamentals of Veterinary Ophthalmology, Elsevier Saunders. [6th ed.]."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1002\/vetr.70618","article-title":"Concern over Crufts \u2018best in show\u2019 winner","volume":"198","author":"Loeb","year":"2026","journal-title":"Vet. Rec."},{"key":"ref_37","first-page":"1","article-title":"Long-term success rate of entropion surgery is equivalent in juvenile and adult dogs","volume":"262","author":"Curto","year":"2023","journal-title":"J. Am. Vet. Med. Assoc."},{"key":"ref_38","doi-asserted-by":"crossref","unstructured":"O\u2019Neill, D.G., Mustikka, M.P., Brodbelt, D.C., Church, D.B., and Vaattovaara, V. (2025). Conformational eyelid disorders in dogs under primary veterinary care in the UK: Epidemiology and clinical management. PLoS ONE, 20.","DOI":"10.1371\/journal.pone.0326526"},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1111\/j.1365-3164.2009.00799.x","article-title":"Hereditary cutaneous mucinosis in shar pei dogs is associated with increased hyaluronan synthase-2 mRNA transcription by cultured dermal fibroblasts","volume":"20","author":"Zanna","year":"2009","journal-title":"Vet. Dermatol."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"10553","DOI":"10.1038\/s41598-022-14483-5","article-title":"Ironing out the wrinkles and folds in the epidemiology of skin fold dermatitis in dog breeds in the UK","volume":"12","author":"Rowe","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_41","first-page":"314","article-title":"Myotonia in the goat","volume":"317","author":"Bryant","year":"1979","journal-title":"Ann. N. Y. Acad. Sci."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"3229","DOI":"10.1002\/ece3.3881","article-title":"Patterns of musculoskeletal growth and dimensional changes associated with selection and developmental plasticity in domestic and wild strain turkeys","volume":"8","author":"Stover","year":"2018","journal-title":"Ecol. Evol."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"EFSA Panel on Animal Health and Welfare (2026). Welfare assessment of turkeys (Meleagris gallopavo gallopavo) on farm. EFSA J., 24, e9851.","DOI":"10.2903\/j.efsa.2026.9851"},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Knowles, T.G., Kestin, S.C., Haslam, S.M., Brown, S.N., Green, L.E., Butterworth, A., Pope, S.J., Pfeiffer, D., and Nicol, C.J. (2008). Leg disorders in broiler chickens: Prevalence, risk factors and prevention. PLoS ONE, 3.","DOI":"10.1371\/journal.pone.0001545"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"2970","DOI":"10.3382\/ps.2014-04291","article-title":"Growth, efficiency, and yield of commercial broilers from 1957, 1978, and 2005","volume":"93","author":"Zuidhof","year":"2014","journal-title":"Poult. Sci."},{"key":"ref_46","unstructured":"Universities Federation for Animal Welfare (UFAW) (2026, May 06). Rolling and Tumbling in Pigeons. Available online: https:\/\/www.ufaw.org.uk\/birds\/pigeons-rolling-and-tumbling."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"10","DOI":"10.1186\/s40575-015-0023-8","article-title":"Epidemiological associations between brachycephaly and upper respiratory tract disorders in dogs attending veterinary practices in England","volume":"2","author":"Jackson","year":"2015","journal-title":"Canine Genet. Epidemiol."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"191","DOI":"10.7120\/09627286.26.2.191","article-title":"Purchasing popular purebreds: Investigating the influence of breed-type on the pre-purchase attitudes and behaviour of dog owners","volume":"26","author":"Packer","year":"2017","journal-title":"Anim. Welf."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Packer, R.M.A., and O\u2019Neill, D.G. (2021). A historical perspective on brachycephalic breed health and the role of the veterinary profession. Health and Welfare of Brachycephalic (Flat-Faced) Companion Animals, CRC Press.","DOI":"10.1201\/9780429263231"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"125","DOI":"10.2307\/4444260","article-title":"Nothing in biology makes sense except in the light of evolution","volume":"35","author":"Dobzhansky","year":"1973","journal-title":"Am. Biol. Teach."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1111\/1365-2745.12547","article-title":"Demography beyond the population","volume":"104","author":"Griffith","year":"2016","journal-title":"J. Ecol."},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"O\u2019Neill, D.G., McMillan, K.M., Church, D.B., and Brodbelt, D.C. (2023). Dog breeds and conformations in the UK in 2019: VetCompass canine demography and some consequent welfare implications. PLoS ONE, 18.","DOI":"10.1371\/journal.pone.0288081"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"100028","DOI":"10.1016\/j.health.2022.100028","article-title":"Longitudinal analysis of the determinants of life expectancy and healthy life expectancy: A causal approach","volume":"2","author":"Aanegola","year":"2022","journal-title":"Healthc. Anal."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"6415","DOI":"10.1038\/s41598-022-10341-6","article-title":"Life tables of annual life expectancy and mortality for companion dogs in the United Kingdom","volume":"12","author":"Teng","year":"2022","journal-title":"Sci. Rep."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1186\/s40575-018-0057-9","article-title":"Demography and disorders of the French Bulldog population under primary veterinary care in the UK in 2013","volume":"5","author":"Baral","year":"2018","journal-title":"Canine Genet. Epidemiol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"81","DOI":"10.7120\/096272812X13345905673809","article-title":"Do dog owners perceive the clinical signs related to conformational inherited disorders as normal for the breed? A potential constraint to improving canine welfare","volume":"21","author":"Packer","year":"2012","journal-title":"Anim. Welf."},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Mellor, D.J., Beausoleil, N.J., Littlewood, K.E., McLean, A.N., McGreevy, P.D., Jones, B., and Wilkins, C. (2020). The 2020 Five Domains Model: Including human-animal interactions in assessments of animal welfare. Animals, 10.","DOI":"10.3390\/ani10101870"},{"key":"ref_58","unstructured":"Farm Animal Welfare Council (FAWC) (1993). Second Report on Priorities for Research and Development in Farm Animal Welfare."},{"key":"ref_59","unstructured":"Brambell, F.W.R. (1965). Report of the Technical Committee to Enquire into the Welfare of Animals Kept Under Intensive Livestock Husbandry Systems (Brambell Report), HMSO."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"243","DOI":"10.1111\/vsu.12735","article-title":"Effects of manipulations to detect sliding hiatal hernia in dogs with brachycephalic airway obstructive syndrome","volume":"47","author":"Broux","year":"2018","journal-title":"Vet. Surg."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"703","DOI":"10.1111\/jsap.12734","article-title":"Documenting the prevalence of hiatal hernia and oesophageal abnormalities in brachycephalic dogs using fluoroscopy","volume":"58","author":"Reeve","year":"2017","journal-title":"J. Small Anim. Pract."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"273","DOI":"10.1111\/j.1748-5827.2005.tb00320.x","article-title":"Prevalence of gastrointestinal tract lesions in 73 brachycephalic dogs with upper respiratory syndrome","volume":"46","author":"Poncet","year":"2005","journal-title":"J. Small Anim. Pract."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"670","DOI":"10.1111\/jsap.12914","article-title":"Relationship between brachycephalic airway syndrome and gastrointestinal signs in three breeds of dog","volume":"59","author":"Kaye","year":"2018","journal-title":"J. Small Anim. Pract."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1186\/s40575-016-0035-z","article-title":"Demography and health of Pugs under primary veterinary care in England","volume":"3","author":"Darwent","year":"2016","journal-title":"Canine Genet. Epidemiol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"521","DOI":"10.1111\/jsap.13325","article-title":"Frequency, breed predisposition and demographic risk factors for overweight status in dogs in the UK","volume":"62","author":"Pegram","year":"2021","journal-title":"J. Small Anim. Pract."},{"key":"ref_66","unstructured":"Royal Kennel Club (2025, January 03). Breed Standards. Available online: https:\/\/www.thekennelclub.org.uk\/breed-standards\/."},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Fawcett, A., Barrs, V., Awad, M., Child, G., Brunel, L., Mooney, E., Martinez-Taboada, F., McDonald, B., and McGreevy, P. (2019). Consequences and management of canine brachycephaly in veterinary practice: Perspectives from Australian veterinarians and veterinary specialists. Animals, 9.","DOI":"10.3390\/ani9010003"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Packer, R.M.A., and O\u2019Neill, D.G. (2021). Dental and oral health for the brachycephalic companion animal. Health and Welfare of Brachycephalic (Flat-Faced) Companion Animals, CRC Press.","DOI":"10.1201\/9780429263231"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1160","DOI":"10.2460\/javma.251.10.1160","article-title":"Effect of brachycephaly and body condition score on respiratory thermoregulation of healthy dogs","volume":"251","author":"Davis","year":"2017","journal-title":"J. Am. Vet. Med. Assoc."},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Gallman, J., Lee-Fowler, T., Clark-Price, S., and Grobman, M. (2023). Evaluation of infrared thermography and 6-minute walk tests to assess airflow limitation, impaired thermoregulation, and exercise intolerance in dogs with brachycephalic obstructive airway syndrome. PLoS ONE, 18.","DOI":"10.1371\/journal.pone.0283807"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"9128","DOI":"10.1038\/s41598-020-66015-8","article-title":"Incidence and risk factors for heat-related illness (heatstroke) in UK dogs under primary veterinary care in 2016","volume":"10","author":"Hall","year":"2020","journal-title":"Sci. Rep."},{"key":"ref_72","doi-asserted-by":"crossref","unstructured":"Hall, E.J., Carter, A.J., and O\u2019Neill, D.G. (2020). Dogs don\u2019t die just in hot cars: Exertional heat-related illness (heatstroke) is a greater threat to UK dogs. Animals, 10.","DOI":"10.3390\/ani10081324"},{"key":"ref_73","doi-asserted-by":"crossref","unstructured":"Packer, R.M.A., O\u2019Neill, D.G., Fletcher, F., and Farnworth, M. (2020). Come for the looks, stay for the personality? A mixed methods investigation of reacquisition and owner recommendation of Bulldogs, French Bulldogs and Pugs. PLoS ONE, 15.","DOI":"10.1371\/journal.pone.0237276"},{"key":"ref_74","doi-asserted-by":"crossref","unstructured":"Roberts, T., McGreevy, P., and Valenzuela, M. (2010). Human induced rotation and reorganization of the brain of domestic dogs. PLoS ONE, 5.","DOI":"10.1371\/journal.pone.0011946"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"131","DOI":"10.1111\/j.1748-5827.2006.00056.x","article-title":"Laryngeal collapse in seven brachycephalic puppies","volume":"47","author":"Pink","year":"2006","journal-title":"J. Small Anim. Pract."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1186\/s40575-017-0045-5","article-title":"Corneal ulcerative disease in dogs under primary veterinary care in England: Epidemiology and clinical management","volume":"4","author":"Lee","year":"2017","journal-title":"Canine Genet. Epidemiol."},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"O\u2019Neill, D.G., Yin, Y., Tetas Pont, R., Brodbelt, D.C., Church, D.B., Pegram, C., and Mustikka, M. (2022). Breed and conformational predispositions for prolapsed nictitating membrane gland (PNMG) in dogs in the UK: A VetCompass study. PLoS ONE, 17.","DOI":"10.1371\/journal.pone.0260538"},{"key":"ref_78","doi-asserted-by":"crossref","unstructured":"Nitzsche, B., Schulze, S., Boltze, J., and Schmidt, M.J. (2023). Reduced cingulate gyrus volume in Cavalier King Charles Spaniels with syringomyelia and neuropathic pain revealed by voxel-based morphometry: A pilot study. Front. Neuroanat., 17.","DOI":"10.3389\/fnana.2023.1175953"},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Knowler, S.P., Cross, C., Griffiths, S., McFadyen, A.K., Jovanovik, J., Tauro, A., Kibar, Z., Driver, C.J., La Ragione, R.M., and Rusbridge, C. (2017). Use of morphometric mapping to characterise symptomatic Chiari-like malformation, secondary syringomyelia and associated brachycephaly in the Cavalier King Charles Spaniel. PLoS ONE, 12.","DOI":"10.1371\/journal.pone.0170315"},{"key":"ref_80","doi-asserted-by":"crossref","unstructured":"McGreevy, P.D., Georgevsky, D., Carrasco, J., Valenzuela, M., Duffy, D.L., and Serpell, J.A. (2013). Dog behavior co-varies with height, bodyweight and skull shape. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0080529"},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1159\/000073756","article-title":"A strong correlation exists between the distribution of retinal ganglion cells and nose length in the dog","volume":"63","author":"McGreevy","year":"2004","journal-title":"Brain Behav. Evol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"14284","DOI":"10.1038\/s41598-023-41229-8","article-title":"The difference between two brachycephalic and one mesocephalic dog breeds\u2019 problem-solving performance suggests evidence for paedomorphism in behaviour","volume":"13","author":"Ujfalussy","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"106095","DOI":"10.1016\/j.tvjl.2024.106095","article-title":"Vertebral and spinal malformations in small brachycephalic dog breeds: Current knowledge and remaining questions","volume":"304","author":"Rohdin","year":"2024","journal-title":"Vet. J."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1163\/156853908783402894","article-title":"Behavioural responses of Canis familiaris to different tail lengths of a remotely-controlled life-size dog replica","volume":"145","author":"Leaver","year":"2008","journal-title":"Behaviour"},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"863","DOI":"10.1111\/ahe.12729","article-title":"Comparative morphometric study of the mimic facial muscles of brachycephalic and dolichocephalic dogs","volume":"50","author":"Schatz","year":"2021","journal-title":"Anat. Histol. Embryol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"106134","DOI":"10.1016\/j.applanim.2023.106134","article-title":"Can my human read my flat face? The curious case of understanding the contextual cues of extremely brachycephalic dogs","volume":"270","author":"Liu","year":"2024","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"e39","DOI":"10.1017\/awf.2024.44","article-title":"Animal affect, welfare and the Bayesian brain","volume":"33","author":"Lecorps","year":"2024","journal-title":"Anim. Welf."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1080\/00480169.2014.940410","article-title":"Introducing breathlessness as a significant animal welfare issue","volume":"63","author":"Beausoleil","year":"2015","journal-title":"N. Z. Vet. J."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"e135","DOI":"10.1002\/vetr.135","article-title":"Owner reported breathing scores, accelerometry and sleep disturbances in brachycephalic and control dogs: A pilot study","volume":"189","author":"Barker","year":"2021","journal-title":"Vet. Rec."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1016\/S0079-6123(08)60069-2","article-title":"Sleep as a restorative process: Human clues","volume":"Volume 53","author":"McConnell","year":"1980","journal-title":"Progress in Brain Research"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"6828","DOI":"10.1038\/s41598-021-86235-w","article-title":"Proposing the VetCompass clinical grading tool for heat-related illness in dogs","volume":"11","author":"Hall","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_92","doi-asserted-by":"crossref","unstructured":"Hall, E.J., Carter, A.J., Chico, G., Bradbury, J., Gentle, L.K., Barfield, D., and O\u2019Neill, D.G. (2022). Risk factors for severe and fatal heat-related illness in UK dogs: A VetCompass study. Vet. Sci., 9.","DOI":"10.3390\/vetsci9050231"},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Hall, E.J., Carter, A.J., Bradbury, J., Beard, S., Gilbert, S., Barfield, D., and O\u2019Neill, D.G. (2023). Cooling methods used to manage heat-related illness in dogs presented to primary care veterinary practices during 2016\u20132018 in the UK. Vet. Sci., 10.","DOI":"10.3390\/vetsci10070465"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"e4153","DOI":"10.1002\/vetr.4153","article-title":"Epidemiology of heat-related illness in dogs under UK emergency veterinary care in 2022","volume":"194","author":"Beard","year":"2024","journal-title":"Vet. Rec."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"4661","DOI":"10.1038\/s41598-025-34768-9","article-title":"Evaluation and validation of a revised VetCompass clinical grading tool for heat-related illness in dogs","volume":"16","author":"Beard","year":"2026","journal-title":"Sci. Rep."},{"key":"ref_96","doi-asserted-by":"crossref","unstructured":"Ilska, J.J., McDonnell, F., and Ladlow, J.F. (2026). Quantitative genetic analysis of respiratory function and related traits in Bulldogs, French Bulldogs and Pugs. PLoS ONE, 21.","DOI":"10.1371\/journal.pone.0348023"},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"1740","DOI":"10.1111\/jvim.15547","article-title":"Retrospective analysis of oesophageal imaging features in brachycephalic versus non-brachycephalic dogs based on videofluoroscopic swallowing studies","volume":"33","author":"Eivers","year":"2019","journal-title":"J. Vet. Intern. Med."},{"key":"ref_98","doi-asserted-by":"crossref","unstructured":"Bottero, E., Ruggiero, P., Falcioni, D., Raponi, F., Campanile, A., De Cata, G., De Lorenzi, D., Gonella, S., Mussi, E., and Borrelli, A. (2025). Clinical evaluation and systematic classification of endoscopic gastrointestinal findings in 176 French Bulldogs with brachycephalic airway obstructive syndrome. Animals, 15.","DOI":"10.3390\/ani15142137"},{"key":"ref_99","doi-asserted-by":"crossref","unstructured":"Quinn, R., Masters, S., Starling, M., White, P.J., Mills, K., Raubenheimer, D., and McGreevy, P. (2025). Functional significance and welfare implications of chewing in dogs (Canis familiaris). Front. Vet. Sci., 12.","DOI":"10.3389\/fvets.2025.1499933"},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"636","DOI":"10.1111\/jsap.13382","article-title":"Keratoconjunctivitis sicca in dogs under primary veterinary care in the UK: An epidemiological study","volume":"62","author":"Brodbelt","year":"2021","journal-title":"J. Small Anim. Pract."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"1763","DOI":"10.1007\/s00429-023-02706-y","article-title":"Sleep-physiological correlates of brachycephaly in dogs","volume":"228","author":"Iotchev","year":"2023","journal-title":"Brain Struct. Funct."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"409","DOI":"10.1136\/vr.104944","article-title":"Canine dystocia in 50 UK first-opinion emergency care veterinary practices: Clinical management and outcomes","volume":"184","author":"Manson","year":"2019","journal-title":"Vet. Rec."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1136\/vr.104108","article-title":"Canine dystocia in 50 UK first-opinion emergency-care veterinary practices: Prevalence and risk factors","volume":"181","author":"Manson","year":"2017","journal-title":"Vet. Rec."},{"key":"ref_104","doi-asserted-by":"crossref","unstructured":"Teng, K.T., Devleesschauwer, B., de Noordhout, C.M., Bennett, P., McGreevy, P.D., Toribio, J.A.L.M.L., and Dhand, N.K. (2018). Welfare-Adjusted Life Years (WALY): A novel metric of animal welfare that combines the impacts of impaired welfare and abbreviated lifespan. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0202580"},{"key":"ref_105","unstructured":"Fisher, S., and Reason, J. (1988). Allostasis: A new paradigm to explain arousal pathology. Handbook of Life Stress, Cognition and Health, Wiley."},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"2093","DOI":"10.1001\/archinte.1993.00410180039004","article-title":"Stress and the individual: Mechanisms leading to disease","volume":"153","author":"McEwen","year":"1993","journal-title":"Arch. Intern. Med."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"272","DOI":"10.1638\/2016-0070.1","article-title":"Evaluating allostatic load: A new approach to measuring long-term stress in wildlife","volume":"49","author":"Edes","year":"2018","journal-title":"J. Zoo Wildl. Med."},{"key":"ref_108","first-page":"261","article-title":"Allostatic load as a measure of animal stress and health risk","volume":"Volume 10","author":"Wolfe","year":"2023","journal-title":"Fowler\u2019s Zoo and Wild Animal Medicine"},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"88","DOI":"10.1016\/j.psyneuen.2014.04.012","article-title":"Do venepuncture procedures induce cortisol responses? A review, study, and synthesis for stress research","volume":"46","author":"Weckesser","year":"2014","journal-title":"Psychoneuroendocrinology"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"2279","DOI":"10.1002\/brv.70179","article-title":"Stress, needs, and behaviour: Understanding chronic stress in pet domestic dogs (Canis lupus familiaris) from the prism of needs","volume":"101","author":"Tananaeva","year":"2026","journal-title":"Biol. Rev."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"106372","DOI":"10.1016\/j.tvjl.2025.106372","article-title":"Static respiratory compliance in anaesthetised and intubated brachycephalic dogs with and without brachycephalic obstructive airway syndrome","volume":"312","author":"Fitzmaurice","year":"2025","journal-title":"Vet. J."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1242\/jcs.092262","article-title":"Chronic hypoxia compromises repair of DNA double-strand breaks to drive genetic instability","volume":"125","author":"Kumareswaran","year":"2012","journal-title":"J. Cell Sci."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"e70046","DOI":"10.1002\/iid3.70046","article-title":"Interactions between sleep, inflammation, immunity and infections: A narrative review","volume":"12","author":"Feuth","year":"2024","journal-title":"Immun. Inflamm. Dis."},{"key":"ref_114","first-page":"1946","article-title":"Stress hormones: Unveiling the role in accelerated cellular senescence","volume":"16","author":"Qin","year":"2024","journal-title":"Aging Dis."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"1822","DOI":"10.1038\/s41591-019-0675-0","article-title":"Chronic inflammation in the etiology of disease across the life span","volume":"25","author":"Furman","year":"2019","journal-title":"Nat. Med."},{"key":"ref_116","doi-asserted-by":"crossref","unstructured":"Word, K.R., Austin, S.H., and Wingfield, J.C. (2022). Allostasis revisited: A perception, variation, and risk framework. Front. Ecol. Evol., 10.","DOI":"10.3389\/fevo.2022.954708"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"132","DOI":"10.1016\/j.ygcen.2006.02.013","article-title":"Actions of glucocorticoids at a seasonal baseline as compared to stress-related levels in the regulation of periodic life processes","volume":"148","author":"Landys","year":"2006","journal-title":"Gen. Comp. Endocrinol."},{"key":"ref_118","unstructured":"All-Party Parliamentary Group for Animal Welfare (APGAW), and O\u2019Neill, D.G. (2026, March 21). Innate Health Assess. Available online: www.innatehealthassessment.org."},{"key":"ref_119","doi-asserted-by":"crossref","unstructured":"Packer, R.M.A., and O\u2019Neill, D.G. (2021). Ethical challenges of treating brachycephalic dogs. Health and Welfare of Brachycephalic (Flat-Faced) Companion Animals, CRC Press.","DOI":"10.1201\/9780429263231"},{"key":"ref_120","unstructured":"Australian Consumer Law (2026, September 02). Competition and Consumer Act 2010 (Cth), Available online: https:\/\/www.legislation.gov.au\/C2004A00109\/latest\/text."},{"key":"ref_121","unstructured":"King\u2019s Printer Acts of Parliament (2026, September 02). The Animal Welfare (Licensing of Activities Involving Animals) (England) Regulations 2018, Available online: https:\/\/www.legislation.gov.uk\/uksi\/2018\/486\/contents\/made."},{"key":"ref_122","unstructured":"NSW Department of Primary Industries (2026, September 02). Animal Welfare Code of Practice: Breeding Dogs and Cats, Available online: https:\/\/www.dpi.nsw.gov.au\/__data\/assets\/pdf_file\/0013\/1310431\/INT21-114956-Breeding-Code-Document.pdf."},{"key":"ref_123","unstructured":"(2026, September 02). Taylor v Lay [2023] NSWCATCD 328; New South Wales Civil and Administrative Tribunal, Appeal Panel, Available online: https:\/\/www.caselaw.nsw.gov.au\/decision\/18c567ad02ea78bd46546ad7."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"e3897","DOI":"10.1002\/vetr.3897","article-title":"\u2018All I do is fight fires\u2019: Qualitative exploration of UK veterinarians\u2019 attitudes towards and experiences of pre-purchase consultations regarding brachycephalic dogs","volume":"194","author":"Pound","year":"2024","journal-title":"Vet. Rec."},{"key":"ref_125","unstructured":"Veterinary Practitioners Board of New South Wales (2026). Assessing fitness for purpose in companion animals. BoardTalk, Veterinary Practitioners Board of New South Wales (VPB NSW)."},{"key":"ref_126","unstructured":"(2026, September 02). Prevention of Cruelty to Animals Act 1979 (NSW), No. 200 of 1979, Available online: https:\/\/legislation.nsw.gov.au\/view\/html\/inforce\/current\/act-1979-200."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"412","DOI":"10.1136\/vr.i5470","article-title":"One Welfare: A platform for improving human and animal welfare","volume":"179","author":"Pinillos","year":"2016","journal-title":"Vet. Rec."}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/16\/18\/2867\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,11]],"date-time":"2026-09-11T14:45:10Z","timestamp":1789137910000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/16\/18\/2867"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,11]]},"references-count":127,"journal-issue":{"issue":"18","published-online":{"date-parts":[[2026,9]]}},"alternative-id":["ani16182867"],"URL":"https:\/\/doi.org\/10.3390\/ani16182867","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,11]]}}}