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324        Small Animal Clinical Nutrition



                                                                      turbance odors that become airborne if such odors are volatile
        VetBooks.ir  Box 18-1. Genes and Olfaction:                   or from the ground, which could include both individual odors
                                                                      and contact/disturbance odors (Hepper and Wells, 2005).
                    People vs. Dogs.
                                                                        Scents tend to be concentrated at their source. Odors spread
                                                                      and become less concentrated, forming a scent cone. When air
                    Olfactory receptors constitute the largest gene family in verte-
                    brates. Species that have highly developed olfactory senses are  is stagnant around the source of an odor, scent pools can form.
                    referred to as macrosmatic (e.g., dogs), whereas species with a  Factors such as wind, terrain, air temperature, humidity and soil
                    weak sense of smell are termed microsmatic (e.g., people).  temperature can affect the scent cone or scent pool, including
                    There are likely several reasons dogs are much better at odor  where and how far it is dispersed (Jones et al, 2004). Hunters
                    detection than people. The surface area of the canine olfactory  using dogs generally agree that the best scent conditions
                    epithelium is as much as 20 times greater than that of people.  include moderate humidity and moderate ambient tempera-
                    The density of neuronal cells and the number of olfactory recep-  tures (early morning and late afternoon on warm sunny days)
                    tors that are expressed on their surface as well as the size of the  (Holloway, 1961). Wind can be channeled by obstructions or
                    olfactory bulb have to be taken into consideration when compar-  terrain and can rapidly disperse scent in unexpected directions
                    ing the olfactory capabilities of different species. The canine
                    olfactory epithelium can express up to 20 times more olfactory  (Jones et al, 2004).Thus, odors do not disperse in a linear con-
                    receptors than that of people. This contributes to the ability of  tinuous gradient but result in odor fragments or patches.These
                    dogs to detect odorant molecules at a much lower concentra-  pockets of odors vary in concentration and are separated by
                    tion. The binding affinity of odorant molecules for their related  areas of clean air where no odor is present.Thus, dogs are often
                    olfactory receptor is also likely to be an important variable that  exposed to variable and intermittent odor signals. In the face of
                    could explain differences in sensing abilities between people  this complexity, dogs and many other animals are able to deter-
                    and dogs. The range of types of olfactory receptors of dogs is  mine the direction of a trail and find the source of an odor.
                    around 30% larger than for people, which could contribute to  Furthermore, they can be trained to identify specific odors
                    the wider range of odorant molecules that dogs can detect.  (Hepper and Wells, 2005). Human handlers who are aware of
                                                                      factors affecting odor dynamics can improve detection success
                    The Bibliography for Box 18-1 can be found at     by directing the activity of a scent-detection dog accordingly
                    www.markmorris.org.
                                                                      (Table 18-2).
                                                                      Olfactory System
                                                                      The olfactory system consists primarily of the nose, nasophar-
                                                                      ynx (including the vomeronasal organ and olfactory epitheli-
                                                                      um), olfactory and vomeronasal nerves and the olfactory bulb
                                                                      (Figure 18-1). The pulmonary system and oro- and nasophar-
                                                                      ynx facilitate olfaction by moving and directing air over the
                                                                      olfactory epithelium.
                                                                        The olfactory epithelium contains the olfactory receptors.
                                                                      This is where odor molecules interact to stimulate an olfactory
                                                                      sensation. Chemicals that are best detected by the olfactory sys-
                                                                      tem are volatile and both water and lipid soluble. Such mole-
                                                                      cules are readily made airborne and dissolve in the mucus that
                                                                      covers the olfactory epithelium. After this occurs, they bind to
                                                                      specific substances called G-protein-coupled-receptors, which
                                                                      are on the cilia of the olfactory receptor neurons. As a result, a
                                                                      signal transduction of smell is initiated (Jones et al, 2004). Box
                                                                      18-1 discusses gene-based differences between the olfactory
                                                                      capabilities of people and dogs.
                                                                        For olfaction to occur, odorants need to contact the olfactory
                                                                      receptors in the olfactory epithelium. Thus, air from the envi-
                                                                      ronment must pass into the nasal cavity to reach the olfactory
                                                                      epithelium. The course taken by air as it passes through the
                                                                      nasal cavity is difficult to predict because of the complex anato-
                  Figure 18-1. Drawing of a mid-sagittal section of a canine head  my of the nasal cavity. There are distinctly different routes of
                  showing olfactory and upper respiratory anatomy.    airflow during normal inspiration and during sniffing. Sniffing
                                                                      draws considerably more air over the olfactory epithelium as
                  als, and as a result, these materials release odors. When follow-  opposed to normal inspiration. During normal inspiration, air
                  ing scent, dogs may be detecting airborne odors arising direct-  tends to route below the olfactory epithelium, a more efficient
                  ly from air moving past an animal or person, from contact/dis-  route for pulmonary function (Becker and King, 1957).
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