Holding back the genes: limitations of research into canine behavioural genetics
Tóm tắt
Canine behaviours that are both desirable and undesirable to owners have a demonstrable genetic component. Some behaviours are breed-specific, such as the livestock guarding by maremmas and flank sucking seen in Dobermanns. While the identification of genes responsible for common canine diseases is rapidly advancing, those genes underlying behaviours remain elusive. The challenges of accurately defining and measuring behavioural phenotypes remain an obstacle, and the use of variable phenotyping methods has prevented meta-analysis of behavioural studies. International standardised testing protocols and terminology in canine behavioural evaluations should facilitate selection against behavioural disorders in the modern dog and optimise breeding success and performance in working dogs. This review examines the common hurdles faced by researchers of behavioural genetics and the current state of knowledge.
Tài liệu tham khảo
Contribution of the pet care industry to the Australian economy http://www.acac.org.au/ACAC_Report_2010.html
U.S. Pet Ownership & Demographics Sourcebook http://www.avma.org/KB/Resources/Statistics/Pages/Market-research-statistics-US-pet-ownership.aspx
Pet Population 2013. http://www.pfma.org.uk/pet-population
McGreevy PD, Bennett P: Challenges and paradoxes in the companion animal niche. Anim Welf 2010,19(S):11–16.
Coppinger R, Zuccotti J: Kennel enrichment: exercise and socialization of dogs. J Appl Anim Welf Sci 1999,2(4):281–296. 10.1207/s15327604jaws0204_3
Lindblad-Toh K, Wade CM, Mikkelsen TS, Karlsson EK, Jaffe DB, Kamal M, Clamp M, Chang JL, Kulbokas EJ 3rd, Zody MC, Mauceli E, Xie X, Breen M, Wayne RK, Ostrander EA, Ponting CP, Galibert F, Smith DR, DeJong PJ, Kirkness E, Alvarez P, Biagi T, Brockman W, Butler J, Chin CW, Cook A, Cuff J, Daly MJ, DeCaprio D, Gnerre S, et al.: Genome sequence, comparative analysis and haplotype structure of the domestic dog. Nature 2005,438(7069):803–819. 10.1038/nature04338
Sutter NB, Eberle MA, Parker HG: Extensive and breed-specific linkage disequilibrium in Canis familiaris . Genome Res 2004, 14: 2388–2396. 10.1101/gr.3147604
Karlsson EK, Baranowska I, Wade CM, Salmon Hillbertz NH, Zody MC, Anderson N, Biagi TM, Patterson N, Pielberg GR, Kulbokas EJ 3rd, Comstock KE, Keller ET, Mesirov JP, von Euler H, Kämpe O, Hedhammar A, Lander ES, Andersson G, Andersson L, Lindblad-Toh K: Efficient mapping of mendelian traits in dogs through genome-wide association. Nature Genet 2007,39(11):1321–1328. 10.1038/ng.2007.10
Karlsson E, Lindblad-toh K: Leader of the pack: gene mapping in dogs and other model organisms. Nature Rev Genet 2008,9(9):713. 10.1038/nrg2382
Jones P, Chase K, Martin A, Davern P, Ostrander EA, Lark KG: Single-nucleotide-polymorphism-based association mapping of dog stereotypes. Genetics 2008,179(2):1033–1044. 10.1534/genetics.108.087866
Vaysse A, Ratnakumar A, Derrien T, Axelsson E, Rosengren Pielberg G, Sigurdsson S, Fall T, Seppala EH, Hansen MS, Lawley CT, Karlsson EK, Consortium LUPA, Bannasch D, Vilà C, Lohi H, Galibert F, Fredholm M, Häggström J, Hedhammar A, André C, Lindblad-Toh K, Hitte C, Webster MT: Identification of genomic regions associated with phenotypic variation between dog breeds using selection mapping. PLoS Genet 2011,7(10):e1002316. 10.1371/journal.pgen.1002316
Hall NJ, Wynne CD: The canid genome: behavioral geneticists’ best friend? Genes Brain Behav 2012, 11: 889–902.
International Schizophrenia Consortium: Common polygenic variation contributes to risk of schizophrenia and bipolar disorder. Nature 2009, 460: 748–752.
Kelley RB: Sheep Dogs. Their Breeding, Maintenance and Training. Sydney, Australia: Halstead Press; 1942.
Scott JP: Genetics and the development of social behavor in Dogs. Am Zool 1964,4(2):161–168.
Scott JP, Fuller JL: Genetics and the social behavior of the Dog. Chicago: The University of Chicago Press; 1965.
Serpell J, Jagoe JA: Early experience and the development of behaviour. In The Domestic Dog: Its evolution, behaviour and interactions with people. Edited by: Serpell J. Cambridge: Cambridge University Press; 1995:79–102.
Overall KL: That dog is smarter than you know: advances in understanding canine learning, memory, and cognition. Top Companion Anim Med 2011,26(1):2–9. 10.1053/j.tcam.2011.01.001
Champagne FA, Curley JP: Epigenetic mechanisms mediating the long-term effects of maternal care on development. Neurosci Biobehav Rev 2009,33(4):593–600. 10.1016/j.neubiorev.2007.10.009
Overall KL, Dunham AE, Frank D: Frequency of nonspecific clinical signs in dogs with separation anxiety, thunderstorm phobia, and noise phobia, alone or in combination. J Am Vet Med Assoc 2001,219(4):467–473. 10.2460/javma.2001.219.467
Bamberger M, Houpt KA: Signalment factors, comorbidity, and trends in behavior diagnoses in dogs: 1,644 cases (1991–2001). J Am Vet Med Assoc 2006,229(10):1591–1601. 10.2460/javma.229.10.1591
Diederich C, Giffroy J: Behavioural testing in dogs: a review of methodology in search for standardisation☆. App Anim Behav Sci 2006,97(1):51–72. 10.1016/j.applanim.2005.11.018
Tomkins LM, McGreevy PD, Branson NJ: Lack of standardization in reporting motor laterality in the domestic dog (Canis familiaris). J Vet Behav:Clin Appl Res 2010,5(5):235–239. 10.1016/j.jveb.2010.03.002
Svartberg K: A comparison of behaviour in test and in everyday life: evidence of three consistent boldness-related personality traits in dogs. App Anim Behav Sci 2005,91(1–2):103–128.
Saetre P, Strandberg E, Sundgren P-E, Pettersson U, Jazin E, Bergström TF: The genetic contribution to canine personality. Gene Brain Behav 2006, 5: 240–248. 10.1111/j.1601-183X.2005.00155.x
Ruefenacht S, Gebhardt-Henrich S, Miyake T, Gaillard C: A behaviour test on German Shepherd dogs: heritability of seven different traits. App Anim Behav Sci 2002,79(2):113–132. 10.1016/S0168-1591(02)00134-X
Thomson PC, Wilson BJ, Wade CM, Shariflou MR, James JW, Tammen I, Raadsma HW, Nicholas FW: The utility of estimated breeding values for inherited disorders of dogs. Vet J 2010,183(3):243–244. 10.1016/j.tvjl.2009.12.002
Arvelius P, Malm S, Svartberg K, Strandberg E: Measuring herding behavior in Border collie-effect of protocol structure on usefulness for selection. J Vet Behav:Clin Appl Res 2013,8(1):9–18. 10.1016/j.jveb.2012.04.007
Hoffmann U, Hamann H, Distl O: Genetic analysis of markers of the performance test for herding dogs. 1. Performance traits. Berl Munch Tierarztl Wochenschr 2003, 116: 81–89.
Karjalainen L, Ojala M, Vilva V: Environmental effects and genetic parameters for measurements of hunting performance in the Finnish Spitz. J Anim Breed Genet 1996, 113: 525–534. 10.1111/j.1439-0388.1996.tb00641.x
Liinamo AE, Karjalainen L, Ojala M, Vilva V: Estimates of genetic parameters and environmental effects for measures of hunting performance in Finnish hounds. J Anim Sci 1997, 75: 622–629.
Hsu Y, Serpell JA: Development and validation of a questionnaire for measuring behavior and temperament traits in pet dogs. J Am Vet Med Assoc 2003,223(9):1293–1300. 10.2460/javma.2003.223.1293
Duffy DL, Hsu Y, Serpell JA: Breed differences in canine aggression. App Anim Behav Sci 2008,114(3–4):441–460.
van den Berg SM, Heuven HCM, van den Berg L, Duffy DL, Serpell JA: Evaluation of the C-BARQ as a measure of stranger-directed aggression in three common dog breeds. App Anim Behav Sci 2010,124(3–4):136–141.
van den Berg L, Schilder MBH, de Vries PAJ, Leegwater PAJ, van Oost BA: Phenotyping of aggressive behavior in golden retriever dogs with a questionnaire. Behav Genet 2006,36(6):882–902. 10.1007/s10519-006-9089-0
Duffy D, Serpell J: Behavioral assessment of guide and service dogs. J Vet Behav:Clin Appl Res 2008,3(4):186–188.
Ley JM, McGreevy P, Bennett PC: Inter-rater and test-retest reliability of the Monash Canine Personality Questionnaire-Revised (MCPQ-R). App Anim Behav Sci 2009,119(1–2):85–90.
Overall KL, Hamilton SP, Chang ML: Understanding the genetic basis of canine anxiety: phenotyping dogs for behavioral, neurochemical, and genetic assessment. J Vet Behav:Clin Appl Res 2006,1(3):124–141. 10.1016/j.jveb.2006.09.004
Overall KL, Crowell-Davis SL, Dehasse J, Fatjó J, Houpt KA, Mills DS, Palestrini C, Beata C, Seksel K: Terminology think tank. J Vet Behav:Clin Appl Res 2006,1(1, 2, 3):29–46. 80–83, 121–123
Jacobs J, Coe J, Niel L: Resource guarding in dogs: achieving expert consensus on definitions, concepts and treatment methods. In 9th International Veterinary Behaviour Meeting: 2013. Lisbon, Portugal: PsiAnimal; 2013:122–123.
McGreevy PD, Calnon D: Getting canine aggression in perspective. Vet J 2010,186(1):1–2. 10.1016/j.tvjl.2009.09.007
Yeh M, Coccaro E, Jacobson K: Multivariate behavior genetic analyses of aggressive behavior subtypes. Behav Genet 2010,40(5):603–617. 10.1007/s10519-010-9363-z
Houpt KA, Willis MB: Genetics of Behavior. In The genetics of the dog. Oxon, New York: CABI Publishing; 2001:371–400.
Ohl F, Arndt SS, van der Staay FJ: Pathological anxiety in animals. Vet J 2008,175(1):18–26. 10.1016/j.tvjl.2006.12.013
Landsberg GM, Hunthausen W, Ackerman L: Fears and phobias. In Handbook of behavior problems of the dog and cat. 2nd edition. Edinburgh, New York: Elsevier Saunders; 2008:227–268.
Mills DS: Medical paradigms for the study of problem behaviour: a critical review. App Anim Behav Sci 2003,81(3):265–277. 10.1016/S0168-1591(02)00286-1
Landsberg GM, Hunthausen W, Ackerman L: Behavior counseling and the veterinary practitioner. In Handbook of behavior problems of the dog and cat. 2nd edition. Edinburgh, New York: Elsevier; 2008:1–14.
Mikkelsen J, Damkjer Lund J: Euthanasia of dogs due to behavioural problems: an epidemiological study of euthanasia of dogs in Denmark, with a special focus on problems of aggression. Eur J Companion Anim Pract 2000,10(2):143–150.
O’Neill DG, Church DB, McGreevy PD, Thomson PC, Brodbelt DC: Longevity and mortality of owned dogs in England. Vet J 2013,198(3):638–643. 10.1016/j.tvjl.2013.09.020
Salmon MD, Hutchison J, Ruch-Gallie R, Kogan L, New JC, Kass PH, Scarlett JM: Behavioral reasons for relinquishment of dogs and cats to 12 shelters. J Appl Anim Welf Sci 2000,3(2):93–106. 10.1207/S15327604JAWS0302_2
Reisner IR: The pathophysiologic basis of behavior problems. Vet Clin North Am Small Anim Pract 1991,21(2):207–224.
Overall KL: Medical differentials with potential behavioral manifestations. Vet Clin North Am Small Anim Pract 2003,33(2):213–229. 10.1016/S0195-5616(02)00103-1
Buxbaum JD, Daly MJ, Devlin B, Lehner T, Roeder K, State MW: The autism sequencing consortium: large-scale, high-throughput sequencing in autism spectrum disorders. Neuron 2012,76(6):1052–1056. 10.1016/j.neuron.2012.12.008
Weber KL, Drake DJ, Taylor JF, Garrick DJ, Kuehn LA, Thallman RM, Schanabel RD, Snelling WM, Pollak EJ, Van Eenennaam AL: The accuracies of DNA-based estimates of genetic merit derived from Angus or multibreed beef cattle training populations. J Anim Sci 2012,90(12):4191–4202. 10.2527/jas.2011-5020
Ollivier L: European pig genetic diversity: a minireview. Animal 2009,3(7):915–924. 10.1017/S1751731109004297
Bilgili SF, Curtis PA: Poultry meat and egg quality symposium introduction. J Appl Poult Res 2007, 16: 91. 10.1093/japr/16.1.91
Wilson BJ, Wade CM: Empowering international canine inherited disorder management. Mamm Genome 2012,23(1–2):195–202.
Hedhammar AA, Malm S, Bonnett B: International and collaborative strategies to enhance genetic health in purebred dogs. Vet J 2011,189(2):189–196. 10.1016/j.tvjl.2011.06.018
Spady TC, Ostrander EA: Canine behavioral genetics: pointing out the phenotypes and herding up the genes. Am J Hum Genet 2008,82(1):10–18. 10.1016/j.ajhg.2007.12.001
Svartberg K: Breed-typical behaviour in dogs—historical remnants or recent constructs? App Anim Behav Sci 2006,96(3–4):293–313.
Mellersh CS: Genetics of Eye Disorders in the Dog. In The Genetics of the Dog, 2nd ed. 2nd edition. Edited by: Ostrander E, Ruvinsky A. Cambridge: CABI; 2012:218–240.
Takeuchi Y, Hashizume C, Arata S, Inoue-Murayama M, Maki T, Hart BL, Mori Y: An approach to canine behavioural genetics employing guide dogs for the blind. Anim Genet 2009,40(2):217–224. 10.1111/j.1365-2052.2008.01823.x
Takeuchi Y, Kaneko F, Hashizume C, Masuda K, Ogata N, Maki T, Inoue-Murayama M, Hart BL, Mori Y: Association analysis between canine behavioural traits and genetic polymorphisms in the Shiba Inu breed. Anim Genet 2009,40(5):616–622. 10.1111/j.1365-2052.2009.01888.x
Dodman NH, Branson NJ, Gliatto J: Tail chasing in a bull terrier. J Am Vet Med Assoc 1993,202(5):758–760.
Moon-Fanelli AA, Dodman NH, Cottam N: Blanket and flank sucking in Doberman Pinschers. J Am Vet Med Assoc 2007,231(6):907–912. 10.2460/javma.231.6.907
Dodman NH, Karlsson EK, Moon-Fanelli A, Galdzicka M, Perloski M, Shuster L, Lindblad-Toh K, Ginns EI: A canine chromosome 7 locus confers compulsive disorder susceptibility. Mol Psychiatr 2010,15(1):8–10. 10.1038/mp.2009.111
Tiira K, Hakosalo O, Kareinen L, Thomas A, Hielm-Bjorkman A, Escriou C, Arnold P, Lohi H: Environmental effects on compulsive tail chasing in dogs. PLoS ONE 2012, 7: e41684. 2012/07/31 edn 10.1371/journal.pone.0041684
Sundqvist AK, Bjornerfeldt S, Leonard JA, Hailer F, Hedhammar A, Ellegren H, Vila C: Unequal contribution of sexes in the origin of dog breeds. Genet 2006,172(2):1121–1128.
Carrasco JJ, Georgevsky D, Valenzuela M, McGreevy PD: A pilot study of sexual dimorphism in the head morphology of domestic dogs. J Vet Behav:Clin Appl Res 2014,9(1):43–46. 10.1016/j.jveb.2013.09.004
Gácsi M, McGreevy P, Kara E, Miklósi A: Effects of selection for cooperation and attention in dogs. Behav Brain Funct 2009, 5: 31. 2009/07/28 edn 10.1186/1744-9081-5-31
McGreevy PD, Georgevsky D, Carrasco JJ, Valenzuela M, Duffy DL, Serpell JA: Dog behavior co-varies with height, bodyweight and skull shape. PLoS 2013,8(12):e80529. 10.1371/journal.pone.0080529
Martínez ÁG, Santamarina Pernas G, Diéguez Casalta FJ, Suárez Rey ML, De la Cruz Palomino LF: Risk factors associated with behavioral problems in dogs. J Vet Behav:Clin Appl Res 2011,6(4):225–231. 10.1016/j.jveb.2011.01.006
Ley JM, Bennett PC, Coleman GJ: A refinement and validation of the Monash Canine Personality Questionnaire (MCPQ). App Anim Behav Sci 2009,116(2–4):220–227.
Baxter T, Hayward J, Castelhano M, Oliveira K, Todhunter R, Serpell J, AR B: Genomic approaches to identify putative canine behavior-associated genes. In Proceedings of the 7th International Conference on Advances in Canine and Feline Genomics and Inherited Diseases: 23–27 September 2013. Cambridge, Massachusetts: The Broad Institute; 2013:38.
Arhant C, Bubna-Littitz H, Bartels A, Futschik A, Troxler J: Behaviour of smaller and larger dogs: effects of training methods, inconsistency of owner behaviour and level of engagement in activities with the dog. App Anim Behav Sci 2010,123(3–4):131–142.
Goddard ME, Beilharz RG: Genetic and environmental factors affecting the suitability of dogs as guide dogs for the blind. Theor Appl Genet 1982,62(2):97–102. 10.1007/BF00293339
Ennik I, Liinamo A-E, Leighton E, van Arendonk J: Suitability for field service in 4 breeds of guide dogs. J Vet Behav:Clin Appl Res 2006,1(2):67–74. 10.1016/j.jveb.2006.06.004
Maejima M, Inoue-Murayama M, Tonosaki K, Matsuura N, Kato S, Saito Y, Weiss A, Murayama Y, Ito S: Traits and genotypes may predict the successful training of drug detection dogs. App Anim Behav Sci 2007,107(3–4):287–298.
Evans RI, Herbold JR, Bradshaw BS, Moore GE: Causes for discharge of military working dogs from service: 268 cases (2000–2004). J Am Vet Med Assoc 2007,231(8):1215–1220. 10.2460/javma.231.8.1215
Wilsson E, Sundgren PE: Behaviour test for eight-week old puppies – heritabilities of tested behaviour traits and its correspondence to later behaviour. App Anim Behav Sci 1998, 58: 151–162. 10.1016/S0168-1591(97)00093-2
Svartberg K: Shyness-boldness predicts performance in working dogs. App Anim Behav Sci 2002, 79: 157–174. 10.1016/S0168-1591(02)00120-X
Wilsson E, Sundgren PE: The use of a behaviour test for selection of dogs for service and breeding. II. Heritability for tested parameters and effect of selection based on service dog characteristics. App Anim Behav Sci 1997, 54: 235–241. 10.1016/S0168-1591(96)01175-6
Goddard ME, Beilharz RG: Genetics of traits which determine the suitability of dogs as guide-dogs for the blind. Appl Anim Ethol 1983,9(3–4):299–315.
van der Waaij EH, Wilsson E, Strandberg E: Genetic analysis of results of a Swedish behavior test on German Shepherd Dogs and Labrador Retrievers. J Anim Sci 2008,86(11):2853–2861. 10.2527/jas.2007-0616
Lindberg S, Strandberg E, Swenson L: Genetic analysis of hunting behaviour in Swedish Flatcoated Retrievers. App Anim Behav Sci 2004,88(3–4):289–298.
Lin L, Faraco J, Li R, Kadotani H, Rogers W, Lin X, Qiu X, de Jong PJ, Nishino S, Mignot E: The sleep disorder canine narcolepsy is caused by a mutation in the Hypocretin ( Orexin ) Receptor 2 gene. Cell 1999, 98: 365–376. 10.1016/S0092-8674(00)81965-0
Dykman RA, Murphree OD, Ackerman PT: Litter patterns in the offspring of nervous and stable dogs: II. Autonomic and motor conditioning. J Nerv Ment Dis 1966,141(4):419–432.
Dykman RA, Murphree OD, Reese WG: Familial anthropophobia in pointer dogs? Arch Gen Psychiatry 1979,36(9):988–993. 10.1001/archpsyc.1979.01780090074008
Murphree OD: Inheritance of human avoidance and inactivity in two strains of the pointer dog. Biol Psychiat 1973,7(1):23–29.
Uhde TW, Malloy LC, Slate SO: Fearful behavior, body size, and serum IFG-1Levels in nervous and normal pointer dogs. Pharmacol Biochem Behav 1992, 43: 263–269.
Klein E, Steinberg SA, Weiss SRB, Matthews DM, Uhde TW: The relationship between genetic deafness and fear-related behaviors in nervous pointer dogs. Physiol Behav 1988, 43: 307–312. 10.1016/0031-9384(88)90192-8
Arvelius P, Klemetsdal G: How Swedish breeders can substantially increase the genetic gain for the English Setter’s hunting traits. J Anim Breed Genet 2013,130(2):142–153. 10.1111/jbg.12026
Riva J, Bondiolotti G, Michelazzi M, Verga M, Carenzi C: Anxiety related behavioural disorders and neurotransmitters in dogs. App Anim Behav Sci 2008,114(1–2):168–181.
Vermeire S, Audenaert K, Dobbeleir A, De Meester R, De Vos F, Peremans K: Evaluation of the brain 5-HT2A receptor binding index in dogs with anxiety disorders, measured with 123 I-5I-R91150 and SPECT. J Nucl Med 2009,50(2):284–289. 10.2967/jnumed.108.055731
Vermeire S, Audenaert K, Vandermeulen E, De Meester R, van Bree H, Dobbeleir A, Peremans K: What’s in a brain: neuroanatomy and neurochemistry of anxiety disorders in dogs. Vlaams Diergeneeskd Tijdschr 2011,80(3):175–184.
Reisner IR, Mann JJ, Stanley M, Huang Y, Houpt KA: Comparison of cerebrospinal fluid monoamine metabolite levels in dominant-aggressive and non-aggressive dogs. Brain Res 1996, 714: 57–64. 10.1016/0006-8993(95)01464-0
Amat M, Le Brech S, Camps T, Torrente C, Mariotti VM, Ruiz JL, Manteca X: Differences in serotonin serum concentration between aggressive English cocker spaniels and aggressive dogs of other breeds. J Vet Behav:Clin Appl Res 2013,8(1):19–25. 10.1016/j.jveb.2012.04.003
Vage J, Lingaas F: Single nucleotide polymorphisms (SNPs) in coding regions of canine dopamine- and serotonin-related genes. BMC Genet 2008, 9: 10.
Vage J, Wade C, Biagi T, Fatjo J, Amat M, Lindblad-Toh K, Lingaas F: Association of dopamine- and serotonin-related genes with canine aggression. Genes Brain Behav 2010,9(4):372–378. 10.1111/j.1601-183X.2010.00568.x
Konno A, Inoue-Murayama M, Hasegawa T: Androgen receptor gene polymorphisms are associated with aggression in Japanese Akita Inu. Biol Lett 2011,7(5):658–660. 10.1098/rsbl.2011.0087
Benjamin J, Li L, Patterson C, Greenberg BD, Murphy DL, Hamer DH: Population and familial association between the D4 dopamine receptor gene and measures of Novelty Seeking. Nature Genet 1996,12(1):81–84. 10.1038/ng0196-81
Lee C, Kim C, Shin S, Shin D, Kang J, Park C: The dopamine D4 receptor polymorphism affects the canine fearfulness. Anim Cell Syst 2008,12(2):77–83. 10.1080/19768354.2008.9647158
Hejjas K, Vas J, Topal J, Szantai E, Ronai Z, Szekely A, Kubinyi E, Horvath Z, Sasvari-Szekely M, Miklosi A: Association of polymorphisms in the dopamine D4 receptor gene and the activity-impulsivity endophenotype in dogs. Anim Genet 2007,38(6):629–633. 10.1111/j.1365-2052.2007.01657.x
Hejjas K, Kubinyi E, Ronai Z, Szekely A, Vas J, Miklósi Á, Sasvari-Szekely M, Kereszturi E: Molecular and behavioral analysis of the intron 2 repeat polymorphism in the canine dopamine D4 receptor gene. Gene Brain Behav 2009,8(3):330–336. 10.1111/j.1601-183X.2008.00475.x
Wan M, Hejjas K, Ronai Z, Elek Z, Sasvari-Szekely M, Champagne FA, Miklosi A, Kubinyi E: DRD4 and TH gene polymorphisms are associated with activity, impulsivity and inattention in Siberian Husky dogs. Anim Genet 2013,44(6):717–727. 10.1111/age.12058
Chase K, Jones P, Martin A, Ostrander EA, Lark KG: Genetic mapping of fixed phenotypes: disease frequency as a breed characteristic. J Hered 2009,100(Supplement 1):537–541.
Yokoyama JS: Investigation of complex neuropsychiatric disorders in the domestic dog: genome-wide surveys for loci underlying noise phobia and adult-onset deafness in purpose-bred dogs. San Francisco: University of California; 2010.
van den Berg L, Vos-Loohuis M, Schilder MB, van Oost BA, Hazewinkel HA, Wade CM, Karlsson EK, Lindblad-Toh K, Liinamo AE, Leegwater PA: Evaluation of the serotonergic genes htr1A, htr1B, htr2A, and slc6A4 in aggressive behavior of golden retriever dogs. Behav Genet 2008,38(1):55–66. 10.1007/s10519-007-9179-7
