Canine Olfaction: Scent, Sign, and Situation



Canine olfaction is a rich field of study for the behavioural sciences and neurosciences, and it is rich in interdisciplinary connections. This chapter will explore the neurocognitive and neuroconative bases of olfaction (the neurophysiological foundations of cognition and motivation), and discuss the behavioural, psychological, and semiotic dimensions of scent processing. It will cover the basic psychophysics of olfaction and the methodologies allowing us to explore this sensory modality, as well as the complex cognitive and motivational dimensions of scent. This chapter will open with an overview of the different disciplines involved in the study of canine olfaction. Some basic anatomy and neuroscience will be reviewed, mostly with direct reference to behaviour and associated psychological processes (e.g., cognitive, motivational, and affective systems). For the behavioural aspect of olfaction, a discussion of the contrasting, yet complementary methods of ethology and experimental psychology will be examined. The importance of both field and laboratory research will be highlighted. Olfaction “in context” will also be discussed in reference to zoosemiotics and in order to understand the canine olfactory psychoethology in its most meaningful and functional dimension: processing “signs” (including symptoms as with dogs trained for biomedical applications such as symptom detection). We will conclude with a short commentary on the human-canine sensory symbiosis with sniffer dogs.


Signal Detection Theory Olfactory System Crime Scene Breath Sample Olfactory Cortex 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



Sincere thanks to Alexandra Horowitz for inviting us to participate in this exciting project. Thank you to all the dogs and their owners that participated in our research over the years, the many hundreds of volunteers working in the lab, Honours and graduate students (for a full list, see Simon Gadbois wants to take the opportunity to thank the many mentors who inspired him over the years, “in order of appearance”: Louis Gadbois, Ward O’Neill, Marvin Krank, Werner Honig, Vincent LoLordo, John Fentress, Peter McLeod, William Moger, and Fred Harrington. You have no idea how much you all contributed to shape and focus that mind of mine and force it to always want to synthesize and keep an open mind, and like coyotes, be happy to be a generalist.


  1. Allen, J. J., Bekoff, M., & Crabtree, R. L. (1999). An observational study of coyote (Canis latrans) scent-marking and territoriality in Yellowstone National Park. Ethology, 105, 289–302.Google Scholar
  2. Arons, C. D., & Shoemaker, W. J. (1992). The distribution of catecholamines and beta-endorphin in the brains of three behaviorally distinct breeds of dogs and their F1 hybrids. Brain Research, 594, 31–39.Google Scholar
  3. Bekoff, M. (2001). Observations of scent-marking and discriminating self from others by a domestic dog (Canis familiaris): Tales of displaced yellow snow. Behavioural Processes, 55(2), 75–79.Google Scholar
  4. Berridge, K. C. (2001). Reward learning: Reinforcement, incentives and expectations. In D. L. Medin (Ed.), Psychology of learning and motivation (Vol. 40, pp. 223–278). San Diego: Academic Press.Google Scholar
  5. Berridge, K. C. (2004). Motivation concepts in behavioral neuroscience. Physiology & Behavior 81(2), 179–209.Google Scholar
  6. Berridge, K. C., & Robinson, T. E. (1998). What is the role of dopamine in reward: Hedonics, learning, or incentive salience? Brain Research Reviews, 28(3), 308–367.Google Scholar
  7. Berridge, K. C., Robinson, T. E., & Aldridge, J. W. (2009). Dissecting components of reward: ‘Liking’, ‘wanting’, and learning. Current Opinion in Pharmacology, 9, 65–73.Google Scholar
  8. Betts, T. (1981). Epilepsy: Questions and answers. Nursing Mirror, 153, 6–9.Google Scholar
  9. Bijland, L. R., Bomers, M. K., & Smulders, Y. M. (2013). Smelling the diagnosis. A review on the use of scent in diagnosing disease. The Netherlands Journal of Medicine, 71(6), 300–307.Google Scholar
  10. Blough, D., & Blough, P. (1977). Animal psychophysics. In W. K. Honig & J. E. R. Staddon (Eds.), Handbook of operant behaviour (pp. 514–539). Englewood Cliffs, NJ: Prentice-Hall Inc.Google Scholar
  11. Blough, D. S. (1966). The study of animal sensory processes by operant methods. In W. K. Honig (Ed.), Operant behavior: Areas of research and application (pp. 345–379). New York, NY: Meredith Publishing Company.Google Scholar
  12. Bradbury, J. W., & Vehrencamp, S. L. (2011). Principles of animal communication (2nd ed.). Sunderland, MA: Sinauer Associates Inc.Google Scholar
  13. Brown, S. W., & Goldstein, L. H. (2011). Can seizure-alert dogs predict seizures? Epilepsy Research, 97, 236–242.Google Scholar
  14. Brown, S. W., & Strong, V. (2001). The use of seizure-alert dogs. Seizure, 10, 39–41.Google Scholar
  15. Buck, L. B. (2000). Smell and taste: The chemical senses. In E. R. Kandel, J. H. Shwartz, & T. M. Jessell (Eds.), Principles of neural science (4th ed., pp. 625–647). New York, NY: McGraw-Hill Companies.Google Scholar
  16. Buszewski, B., Ligor, T., Jezierski, T., Wenda-Piesik, A., Walczak, M., & Rudnicka, J. (2012). Identification of volatile lung cancer markers by gas chromatography-mass spectrometry: Comparison with discrimination by canines. Analytical and Bioanalytical Chemistry, 404, 141–146.Google Scholar
  17. Chen, M., Daly, M., & Williams, G. (2000). Non-invasive detection of hypoglycaemia using a novel, fully biocompatible and patient friendly alarm system. British Medical Journal, 321, 1565–1566.Google Scholar
  18. Cornu, J. N., Cancel-Tassin, G., Ondet, V., Girardet, C., & Cussenot, O. (2011). Olfactory detection of prostate cancer by dogs sniffing urine: A step forward in early diagnosis. European Urology, 59, 197–201.Google Scholar
  19. Dalziel, D. J., Uthman, B. M., McGorray, S. P., & Reep, R. L. (2003). Seizure-alert dogs: A review and preliminary study. Seizure, 12, 115–120.Google Scholar
  20. Depue, R. (2000). Neurobehavioral systems, personality and psychopathology. New York, NY: Springer.Google Scholar
  21. Deschênes, M., Moore, J., & Kleinfeld, D. (2011). Sniffing and whisking in rodents. Current Opinion in Neurobiology, 22, 1–8.Google Scholar
  22. Diamond, A. (2006). Bootstrapping conceptual deduction using physical connection: Rethinking frontal cortex. Trends in Cognitive Sciences, 10, 212–218.Google Scholar
  23. Diamond, A., Churchland, A., Cruess, L., & Kirkham, N. (1999). Early developments in the ability to understand the relation between stimulus and reward. Developmental Psychology, 35, 1507–1517.Google Scholar
  24. Ehmann, R., Boedeker, E., Friedrich, U., Sagert, J., Dippon, J., Friedel, G. et al. (2012). Canine scent detection in the diagnosis of lung cancer: Revisiting a puzzling phenomenon. European Respiratory Journal, 39, 669–676.Google Scholar
  25. Engeman, R. M., Vice, D .S., Rodriguez, D. V., Gruver, K. S., Santos, W. S., & Pitzler, M. E. (1998). Effectiveness of the detector dogs used for deterring the dispersal of Brown Tree Snakes. Pacific Conservation Biology, 4, 256–260.Google Scholar
  26. Fentress, J. C., & Gadbois, S. (2001). The development of action sequences. In E. M. Blass (Ed.), Handbooks of behavioral neurobiology: Developmental psychobiology, developmental neurobiology and behavioral ecology: Mechanisms and early principles (Vol. 13, pp. 393–431). New York: Kluwer Academic Publishers.Google Scholar
  27. Fjellanger, R., Andersen, E. K., & McLean, I. G. (2002). A training program for filter-search mine detection dogs. International Journal of Comparative Psychology, 15, 277–286.Google Scholar
  28. Flannery, M., & Gadbois, S. (2013). The use of scent detection dogs in wildlife conservation. Manuscript in preparation.Google Scholar
  29. Furton K. G., & Myers L. J. (2001). The scientific foundations and efficacy of the use of canines as chemical detectors for explosives. Talanta, 54(3), 487–500.Google Scholar
  30. Gadbois, S. (2010). Canine behavioural neuroscience: From canine science in shackles to new opportunities. In Proceedings of the 2nd Canine Science Forum, Vienna, Austria.Google Scholar
  31. Gadbois, S., Demontfaucon, M., Mousse, D., & Flannery, M. (in prep). Ribbon Snake Conservation Canines in Kejimkujik National Park. Google Scholar
  32. Gheusi, G., Goodall, G., & Dantzer, R. (1997). Individually distinctive odours represent individual conspecifics in rats. Animal Behaviour, 53, 935–944.Google Scholar
  33. Gordon, R. T., Schatz, C. B., Myers, L. J., Kosty, M., Gonczy, C., Kroener, J. et al. (2008). The use of canines in the detection of human cancers. The Journal of Alternative and Complementary Medicine, 14, 61–67.Google Scholar
  34. Gotzche, P. C., & Nielsen, M. (2006). Screening for breast cancer with mammography. Cochrane Database of Systematic Reviews, 4, CD001877.Google Scholar
  35. Gray, J. A. (1987). The psychology of fear and stress. New York, NY: Cambridge University Press.Google Scholar
  36. Hall, N. J., Smith, D. W., Wynne, C. D. L. (2013). Training domestic dogs (Canis lupus familiaris) on a novel discrete trials odor-detection task. Learning and Motivation, 44(4), 218–228.Google Scholar
  37. Haberly, L. B. (1998). Olfactory cortex. In G .M. Shepherd (Ed.), The synaptic organization of the brain (4th ed.), (pp. 377–416). New York, NY: Oxford University Press.Google Scholar
  38. Helton, W. S. (2009a). Attention in dogs: Sustained attention in mine detection as case study. In W. S. Helton (Ed.), Canine ergonomics. The science of working dogs (pp. 83–97). Boca Raton, FL: Taylor and Francis Group.Google Scholar
  39. Helton, W. S. (2009b). Overview of scent detection work. In W. S. Helton (Ed.), Canine ergonomics. The science of working dogs (pp. 83–97). Boca Raton, FL: Taylor and Francis Group.Google Scholar
  40. Hepper, P. G., & Wells, D. L. (2005). How many footsteps do dogs need to determine the direction of an odour trail? Chemical Senses, 30, 291–298.Google Scholar
  41. Harrington, F. H., & Asa, C. S. (2003). Wolf communication. In D. Mech & L. Boitani (Eds.), Wolves. Behaviour, ecology, and conservation. (pp. 66–103). Chicago, IL: University of Chicago Press.Google Scholar
  42. Hewes, G. W. (1994). Evolution of human semiosis and the reading of animal tracks. In W. Nöth (Ed.), Origins of semiosis. Sign evolution in nature and culture (pp. 139–149). Berlin, Germany: Walter de Gruyter & Co.Google Scholar
  43. Honig, W. K., & James, P. H. R. (1971). Animal memory. New York, NY: Academic Press.Google Scholar
  44. Honig, W. K. (1978). Studies of working memory in the pigeon. In S. H. Hulse, H. Fowler, & W. K. Honig (Eds.), Cognitive processes in animal behavior (pp. 211–247). Hillsdale, NJ: Lawrence Erlbaum Associates.Google Scholar
  45. Honig, W. K. (1966). Operant behavior: Areas of research and application. New York: Appleton-Century-Crofts.Google Scholar
  46. Honig, W. K. (1981). Working memory and the temporal map. In N. E. Spear & R. R. Miller (Eds.), Information processing in animals: Memory mechanisms (pp. 167–197). Hillsdale, NJ: Lawrence Erlbaum AssociatesGoogle Scholar
  47. Honig, W. K. (1984). Contributions of animal memory to the study of animal learning. In H. L. Roitblat, T. G. Bever, & H. S. Terrace (Eds.), Animal cognition (pp. 29–44). Hillsdale, NJ: Lawrence Erlbaum Associates.Google Scholar
  48. Honig, W. K. & Fetterman, J. G. (1992). Cognitive aspects of stimulus control. Hillsdale, NJ: Lawrence Erlbaum Associates.Google Scholar
  49. Honig, W. K. & Staddon, J. E. R. (1977). Handbook of operant behavior. Englewood Cliffs, NJ: Prentice-Hall.Google Scholar
  50. Horowitz, A., Hecht, J., Dedrick, A. (2013). Smelling more or less: Investigating the olfactory experience of the domestic dog. Learning and motivation, 44, 207–217.Google Scholar
  51. Horvath, G., Järverud, G. K., Järverud, S., & Horváth, I. (2008). Human ovarian carcinomas detected by specific odor. Integrative Cancer Therapy, 7(2), 76–80.Google Scholar
  52. Hulse, S. H., Fowler, H., & Honig, W. K. (1978). Cognitive processes in animal behavior. Hillsdale, NJ: Lawrence Erlbaum Associates.Google Scholar
  53. Jett, J. R. (2005). Limitations of screening for lung cancer with low-dose spiral computer tomography. Clinical Cancer Research, 11, 4988s–4992s.Google Scholar
  54. Kepecs, A., Uchida, N., & Mainen, Z. F. (2005). The sniff as a unit of olfactory processing. Chemical Senses, 31, 167–179.Google Scholar
  55. Kersting, E., Belényi, B., Topál, J., & Miklósi, A. (2009). Judging the effect of epilepsy-seizure alert dogs on human well-being by a self-administered questionnaire. Journal of Veterinary Behavior, 4(2), 84.Google Scholar
  56. Kirton, A., Winter, A., Wirrel, E., & Snead, O. C. (2008). Seizure response dogs: Evaluation of a formal training program. Epilepsy & Behaviour, 13, 499–504.Google Scholar
  57. Kringelbach, M. L., & Berridge, K. C. (2009). Towards a functional neuroanatomy of pleasure and happiness. Trends in Cognitive Sciences, 13, 479–487.Google Scholar
  58. Lit, L. (2009). Evaluating learning tasks commonly applied in detection dog training. In W. S. Helton (Ed.), Canine ergonomics. The science of working dogs (pp. 99–114). Boca Raton, FL: Taylor and Francis Group.Google Scholar
  59. MacLean, P. D. (1990). The triune brain in evolution: Role in paleocerebral functions. New York: Plenum Press.Google Scholar
  60. Macmillan, N. A., & Creelman, C. D. (2005). Detection theory. A user’s guide (2nd ed.). Mahwah, NJ: Lawrence Erlbaum Associates, Inc.Google Scholar
  61. Mainland, J., & Sobel, N. (2006). The sniff is part of the olfactory percept. Chemical Senses, 31, 181–196.Google Scholar
  62. McCulloch, M., Jezierski, T., Broffman, M., Hubbard, A., Turner, K., & Janecki, T. (2006). Diagnostic accuracy of canine scent detection in early- and late-stage lung and breast cancers. Integrative Cancer Therapies, 5(1), 30–39.Google Scholar
  63. McLean, I. G., Bach, H., Fjellanger, R., & Akerblom, C. (2003). Bringing the minefield to the detector: Updating the REST concept. Proceedings of EUDEM2-SCOT, 1, 156–161.Google Scholar
  64. McNicol, D. (2005). A primer of signal detection theory. Mahwah, NJ: Lawrence Erlbaum Associates, Inc.Google Scholar
  65. Menini, A. (2009). The neurobiology of olfaction. Boca Raton, FL: CRC Press.Google Scholar
  66. Miekish, W., Schubert, J. K., & Noeldge-Schomburg, G. F. E. (2004). Diagnostic potential of breath analysis—focus on volatile organic compounds. Clinica Chimica Acta, 347, 25–39.Google Scholar
  67. Moser, E., & McCulloch, M. (2010). Canine scent detection of human cancers: A review of methods and accuracy. Journal of Veterinary Behaviour, 5, 145–152.Google Scholar
  68. Overman, W. H. (1990). Performance on traditional matching to sample, non-matching to sample, and object discrimination tasks by 12–32-month-old children. In A. Diamond (Ed.), The development and neural bases of higher cognitive functions, annals of the New York academy of sciences (Vol. 608, pp. 365–393). New York, NY: New York Academy of Sciences.Google Scholar
  69. Panksepp, J., & Biven, L. (2012). The archaeology of mind: Neuroevolutionary origins of human emotions. New York, NY: W.W. Norton.Google Scholar
  70. Panksepp, J. (1998). Affective neuroscience. The foundations of human and animal emotions. New York, NY: Oxford University Press.Google Scholar
  71. Pastore, R. E., Crawley, E. J., Berens, M. S., & Skelley, M. A. (2003). “Nonparametric” A’ and other modern misconceptions about signal detection theory. Psychonomic Bulletin & Review, 10(3), 556–569.Google Scholar
  72. Pauling, L., Robinson, A. B., Teranishi, R., & Cary, P. (1971). Quantitative analysis of urine vapor and breath by gas–liquid partition chromatography. Proceedings of the National Academy of Science, 68, 2374–2376.Google Scholar
  73. Pearsall, M. D., & Verbruggen, H. (1982). Scent. Training to track, search, and rescue. Loveland, CO: Alpine Publications.Google Scholar
  74. Phillips, M., Herrera, J., Krishnan, S., Zain, M., Greenberg, J., & Cataneo, R. N. (1999). Variation in volatile organic compounds in the breath of normal humans. Journal of Chromatography B, 729, 75–88.Google Scholar
  75. Pickel, D., Manucy, G. P., & Walker, D. B. (2004). Evidence for canine olfactory detection of melanoma. Applied Animal Behaviour Science, 89, 107–116.Google Scholar
  76. Premack, D. (1983). The codes of man and beasts. Behavioral and Brain Sciences, 6(1), 125–137.Google Scholar
  77. Price, J. L. (2003). The olfactory system. In: G. Paxinos (Ed.), The human nervous system (2nd ed)., (pp. 1198–1212). San Diego, CA: Elsevier Academic Press.Google Scholar
  78. Schneider, G. E. (1969). Two visual systems. Science, 163(3870), 895–902.Google Scholar
  79. Schoon, G. A., & Haak, R. (2002). K9 suspect discrimination: Training and practicing scent identification line-ups. Calgary, Alberta: Detselig Enterprises.Google Scholar
  80. Sebeok, T. A. (1968). Animal Communication: Techniques of study and results of research. Bloomington, IN: Indiana University PressGoogle Scholar
  81. Sebeok, T. A. (1977). How animals communicate. Bloomington, IN: Indiana University Press.Google Scholar
  82. Séguinot, V., Cattet, J., & Benhamou, S. (1998). Path integration in dogs. Animal Behaviour, 55, 787–797.Google Scholar
  83. Shepherd, G. M. (1994). Neurobiology (3rd ed.). New York, NY: Oxford University Press.Google Scholar
  84. Slotnick, B., & Schellinck, H. (2002). Methods in olfactory research with rodents. In S. A. Simon & M. Nicolelis (Eds.), Frontiers and methods in chemosenses (pp. 21–61). Boca Raton, FL: CRC Press.Google Scholar
  85. Smith, D. A., Ralls, K., Hurt, A., Adams, B., Parker, M., Davenport, B., et al. (2003). Detection and accuracy rates of dogs trained to find scats of San Joaquin kit foxes (Vulpes macrotis mutica). Animal Conservation, 6, 339–346.Google Scholar
  86. Sobel, N., Prabhakaran, V., Desmond, J. E., Glover, G. H., Goode, R. L., Sulliva, E. V., et al. (1998). Sniffing and smelling: Separate subsystems in the human olfactory cortex. Nature, 392, 282–286.Google Scholar
  87. Sonoda, H., et al. (2011). Colorectal cancer screening with odour material by canine scent detection. Gut, 60, 814–819.Google Scholar
  88. Steen, J. B., & Wilson, E. (1990). How do dogs determine the direction of tracks? Acta Physiologica Scandinavica, 139(4), 531–534Google Scholar
  89. Strong, V., Brown, S., & Walker, R. (1999). Seizure-alert dogs - fact or fiction? Seizure, 8, 62–65.Google Scholar
  90. Strong, V., Brown, S., Huyton, M., & Coyle, H. (2002). Effect of trained Seizure Alert Dogs ® on frequency of tonic-clonic seizures. Seizure, 11, 402–405.Google Scholar
  91. Szulejko, J. R., McCulloch, M., Jackson, J., McKee, D. L., Walker, J. C., & Touradj, S. (2010). Evidence for cancer biomarkers in exhaled breath. IEEE Sensors Journal, 10(1), 185–210Google Scholar
  92. Terrace, H. S. (1963a). Discrimination learning with and without errors. Journal of Experimental Analysis of Behavior, 6, 1–27.Google Scholar
  93. Terrace, H. S. (1963b). Errorless transfer of a discrimination across two continua. Journal of Experimental Analysis of Behavior, 6, 223–232.Google Scholar
  94. Terrace, H. S. (1964). Wavelength generalization after discrimination learning with and without errors. Science, 144, 78–80.Google Scholar
  95. Terrace, H. S. (1966). Stimulus control. In W. K. Honig (Ed.), Operant behavior: Areas of research and application (pp. 271–344). New York: Appleton-Century-Croft.Google Scholar
  96. Thesen, A., Steen, J. B., & Doving, K. B. (1993). Behaviour of dogs during olfactory tracking. Journal of Experimental Biology, 180, 247–251.Google Scholar
  97. Vaché, M., Ferron, J., & Gouat, P. (2001). The ability of Red Squirrels (Tamiasciurus hudsonicus) to discriminate conspecific olfactory signatures. Canadian Journal of Zoology, 79, 1296–1300.Google Scholar
  98. Waggoner, L. P., Jones, M., Williams, M., Johnston, J. M., Edge, C., & Petrousky, J. A. (1998). Effects of extraneous odors on canine detection. SPIE Proceedings, 2575, 355–362.Google Scholar
  99. Wells, D. (2007). Domestic dogs and human health: An overview. British Journal of Health Psychology, 12, 145–156.Google Scholar
  100. Wells, D. L., & Hepper, P. G. (2003). Directional tracking in the domestic dog, Canis familiaris, 84(4), 297–305.Google Scholar
  101. Wells, D. L., Lawson, S. W., & Siriwardena, A. N. (2008). Canine responses to hypoglycemia in patients with Type 1 Diabetes. The Journal of Alternative and Complementary Medicine, 14(10), 1235–1241.Google Scholar
  102. Wells, D. L., Lawson, S. W., & Siriwardena, A. N. (2011). Feline responses to hypoglycemia in people with Type 1 Diabetes. The Journal of Alternative and Complementary Medicine, 17(2), 99–100.Google Scholar
  103. Wells, M. C., & Bekoff, M. (1981). An observational study of scent-marking in coyotes, Canis latrans. Animal Behaviour, 29(2), 332–250.Google Scholar
  104. Williams, H., & Pembroke, A. (1989). Sniffer dogs in the melanoma clinic? Lancet, 333(8640), 734.Google Scholar
  105. Willis, C. M., Church, S. M., Guest, C. M., Cook, W. A., McCarthy, N., Bransbury, A. J., et al. (2004). Olfactory detection of human bladder cancer by dogs: Proof of principle study. British Medical Journal, 329, 712–714.Google Scholar
  106. Wilson, D. A., & Sullivan, R. M. (2011). Cortical Processing of Odor Objects. Neuron 72, 506–519.Google Scholar
  107. Wilson, D. A. & Stevenson, R. J. (2006). Learning to smell: Olfactory perception from neurobiology to behavior. Baltimore: John Hopkins University Press.Google Scholar
  108. Zelano, C., & Sobel, N. (2005). Humans as an Animal Model for Systems-Level Organization of Olfaction. Neuron, 48, 431–454.Google Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2014

Authors and Affiliations

  1. 1.Department of Psychology and Neuroscience, Neuroscience InstituteDalhousie UniversityHalifaxCanada

Personalised recommendations