Elwood, R. W. 2019. Assessing the potential for pain in crustaceans and other invertebrates. In: The Welfare of Invertebrate Animals. Carere, C., Mather, J. (eds), 147-177. Springer, Cham.

All animals face hazards that cause tissue damage, and most have nociceptive reflex responses that protect them from such damage. However, some taxa have also evolved the capacity for pain experience, presumably to enhance long-term protection through behaviour modification based on memory of the unpleasant nature of pain. In this review, I consider various criteria that might help to distinguish nociception from pain. Because nociceptors are so taxonomically widespread, simply demonstrating their presence is not sufficient. Further, investigation of the central nervous system provides limited clues about the potential to experience pain. Opioids and other analgesics might indicate a central modulation of responses, but often, peripheral effects could explain the analgesia. Thus, reduction of responses by analgesics and opioids does not allow clear discrimination between nociception and pain. Physiological changes in response to noxious stimuli or the threat of a noxious stimulus might prove useful, but, to date, application to invertebrates is limited. Behaviour of the organism provides the greatest insights. Rapid avoidance learning and prolonged memory indicate central processing rather than mere reflexes and are consistent with the experience of pain. Complex, prolonged grooming or rubbing may demonstrate an awareness of the specific site of stimulus application. Trade-offs with other motivational systems indicate central processing, and an ability to use complex information suggests sufficient cognitive ability for the animal to have a fitness benefit from a pain experience. Recent evidence of fitness enhancing, anxiety-like states is also consistent with the idea of pain. Thus, available data go beyond the idea of just nociception, but the impossibility of total proof of pain means they are not definitive. Nevertheless, more humane care for invertebrates is suggested.