animal. Training and experience in studying and observing animal behavior are required to interpret what we observe in nonhuman animals.

Nociception

The important distinction between pain and nociception must also be made. Nociception is the term introduced almost 100 years ago by the great physiologist Sherrington (1906) to make clear the distinction between detection of a noxious event or a potentially harmful event and the psychological and other responses to it. Sherrington and others before him understood that pain was not a simple sensation, but rather was a complex experience, only a part of which was sensory in nature. Accordingly, it is most accurate to describe what we study as pain in nonhuman animals as nociception. However, although nonhuman animals cannot express in words the psychological and emotional consequences of a noxious stimulus or event, none of us in this audience would hesitate to apply the term “pain” to that circumstance. This fundamental distinction between pain and nociception emphasizes the importance of interpretation of animal behavior by an experienced individual to assess the presence and intensity of pain and distress.

Having said this, it is nevertheless imperative to acknowledge that unless it is established to the contrary, we should assume that those procedures that produce pain in us might also produce pain in animals. This is an entirely appropriate guideline, bearing in mind the caveats that I discussed and the comments Dr. Bayne made just before me.

Noxious Stimuli

I want to consider briefly the nociceptive apparatus (i.e., basic anatomy and physiology). It is important to do this because our concerns about pain in animals are not related to acute pain associated with procedures that are of short duration and/or simple analgesiometric tests, as Dr. Bayne mentioned, such as a tail flick test or a hot plate test. Rather, we are more concerned about the consequences of procedures (surgical procedures included) that may be associated with longer lasting pain that may cause distress.

In Figure 1, a piece of skin that is innervated by a variety of sensory receptors called nociceptors is illustrated. Nociceptors respond only to stimuli that damage tissue or have the potential to damage tissue, and there are several kinds of nociceptors identified in Figure 1. When a nociceptor is activated by a mechanical, thermal, or chemical stimulus, the stimulus energy is transduced by the nociceptor to an electrical event (action potential) and the information is conveyed along nerve axons to the spinal cord, a part of the central nervous system. Input from nociceptors is transferred in the spinal cord dorsal horn to spinal neuron cell bodies whose axons ascend to supraspinal (brain) sites. Nociceptive information is thus distributed to multiple brain sites that give rise to both simple and complex responses to the peripheral noxious event.



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