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               Making Sense of Scents: Olfactory Perception in Animals
             Neil Jordan




             Olfaction is the action or capacity of smelling   the  secretions  of  such  specialised  glands,
             and is arguably the oldest form of communi-  body  orifices  associated  with  digestion  and
             cation in nature. Olfaction underpins critical   reproduction are also sources of scent signals
             interactions as diverse as chemotaxis in sin-  used in olfactory communication (e.g. Jordan
             gle celled organisms (Vickers 2000), the pol-  et  al. 2013). Whilst chemical information
             lination of flowering plants (Wright and   may be excreted or secreted passively from
             Schiestl 2009), and the nipple‐searching   the bodies of animals as they move through
             behaviour exhibited by newborn mammals   their environment, animals often actively
             (Hudson and Distel 1983). Whilst all forms of   deposit olfactory signals into the environ-
             communication involve the provision of   ment using a diverse range of behaviours col-
             information by a sender and the receipt and   lectively called scent marking.
             response to  that  information  by  a receiver   As scent marks may be costly to produce
             (Bradbury and Vehrencamp 1998), olfaction   and supplies are limited (Gosling et al. 2000),
             contrasts with other forms of communica-  scent marks are adapted and released so as to
             tion in that it does not require close proxim-  maximise their longevity in the environment
             ity between senders and receivers (Bradbury   and the likelihood of their detection by target
             and Vehrencamp 1998). This increases the   receivers (e.g. Roberts and Lowen 1997).
             breadth of contexts in which olfaction may   Both chemical and behavioural adaptions
             be employed and has profound effects on the   may be employed to ensure that signals
             design, distribution, and perception of olfac-  remain in the environment for as long as pos-
             tory signals and the anatomical equipment   sible. For example, fatty acids are commonly
             used in this process.                    found in scent secretions and are thought to
                                                      slow the release of signalling components
                                                      (Alberts 1992), a goal which may be further
               Scent Production and                   promoted by the observation that animals
             Distribution                             commonly  place  their  scents  in  sites  pro-
                                                      tected from the elements (Eisenberg and
             Animals from ants to elephants are covered   Kleiman 1972). Additionally species often
             with an array of exocrine glands. These con-  scent mark in prominent places, such as
             gregations of sebaceous and apocrine tissues   the  crossroads of trails (Barja et  al. 2004),
             are often under the control of hormones,   presumably to promote the discovery of sig-
             which control the production and release of   nals  by  intended  receivers.  The  intended
             chemical signals (Ebling 1977). In addition to   recipients of these olfactory messages vary



             Zoo Animal Learning and Training, First Edition. Edited by Vicky A. Melfi, Nicole R. Dorey, and Samantha J. Ward.
             © 2020 John Wiley & Sons Ltd. Published 2020 by John Wiley & Sons Ltd.
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