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                                   Cerebral cortex      on which it synapses, making it respond
                                Conscious perception    more vigorously to subsequent stimulation.
  VetBooks.ir                       Limbic system       This is called windup or spinal facilitation

                                                        of pain.
                                  Emotional meaning
                                                           Activity in spinal nociceptive pathways
                         Hypothalamus and brainstem
                                  Autonomic reflexes    is also strongly influenced by descending
                                                        antinociceptive systems that originate in
                                                        the brainstem. The midbrain and medulla
                                                        both possess  midline nuclei  that  inhibit
                                                        nociception via their connections with
                                                        nociceptive pathways. These nuclei use a
                                                        variety of neurotransmitters, most notably
                                  PNS   CNS             endorphins, transmitters with powerful
                                                        antinociceptive properties.
                                                           Stoicism, an apparent indifference to
                       Trigeminal n.                    pain, is largely determined by personality
                                                        and training among both humans and
                                                        animals. High‐strung individuals may
                                                        exhibit exaggerated reactions to stimuli
                                 Local reflex arcs      that scarcely merit attention in more
                                                        “laid‐back” individuals. Interestingly,
                        Spinal nn.                      the effectiveness of applying a twitch to a
                                                        horse’s upper lip as a method of restraint
                                                        during mildly painful procedures has
                                                        long been attributed to redirecting the
                                                        horse’s attention from the procedure to
               Figure 12-3.  Divergence in nociceptive path­  the moderately noxious stimulus of the
               ways. The primary afferent neuron of cranial and   twitch. Studies have shown, however,
               spinal nerves brings information about painful   that the correctly applied twitch actually
               stimuli into the central nervous system (CNS).   stimulates a release of endorphins,
               From there the information stimulates protective   lowers the heart rate, and produces
               reflex movements and is sent to other parts of
               the  CNS to produce autonomic and emotional   behavioral signs of sedation in the horse.
               responses. Perception in the cortex is only one   This very old technique of equine restraint
               destination for nociception. n(n.), nerve(s). PNS,   may actually be recruiting some of the
               peripheral nervous system.               neuroanatomic pathways postulated to
                                                        be stimulated by acupuncture.
               of primary hyperalgesia, a phenomenon
               wherein the perception of pain in injured   Proprioception
               tissues is increased. A familiar and dra-
               matic example of this is made by sun-    Proprioception is the nonvisual perception
               burned human skin; the inflammation      of body position. It is a complex sensory
               of the injured skin lowers threshold of   modality that is created through the input
               nociceptors so that even a light touch   of a variety of receptors collectively called
               (for instance, contact with clothing) can   proprioceptors. These include joint recep­
               activate them.                           tors (providing information on tension
                  Events in the dorsal horn of the spinal   and pressure within joints), muscle spindles
               cord can also affect transmission of nocic­  (signaling changes in muscle length), Golgi
               eptive information. Rapid, prolonged firing   tendon organs (signaling tension in tendons),
               of action potentials in the primary afferent   and skin mechanoreceptors (which report
               neuron can produce changes in the neuron   physical contact with the environment).
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