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Helmut Segner Fish Nociception and pain A biological perspective

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hyperalgesia (Tracey 2005). Also, it is at the level of the brainstem<br />

where nociceptive information can be integrated with arousal or<br />

autonomic processes, thereby linking nociception with overall<br />

homeo static regulation in the organism.<br />

The large distributed network of interconnected cortical, subcortical<br />

<strong>and</strong> brainstem regions that is accessed during nociceptive<br />

processing is commonly referred to as <strong>pain</strong> circuit or “<strong>pain</strong> matrix”<br />

(Figures 3 <strong>and</strong> 4). The existence of this matrix highlights that <strong>pain</strong><br />

perception is not a matter of one specific brain region or brain part,<br />

but arises from a concerted network activity. Based on neuroanatomical,<br />

neurophysiological, pharmacological <strong>and</strong> neuroimaging<br />

data, the <strong>pain</strong> matrix is often subdivided into a lateral <strong>and</strong> medial<br />

<strong>pain</strong> systems, with both systems containing brainstem, subcortical<br />

as well as cortical components (Schnakers <strong>and</strong> Zasler 2007,<br />

Tracey <strong>and</strong> Mantyh 2007). The literature is not always consistent<br />

regarding what regions are to be included in the two systems, but<br />

generally the lateral <strong>pain</strong> system is considered to consist of the<br />

lateral thalamus, primary <strong>and</strong> secondary somatosensory cortex (SI<br />

<strong>and</strong> SII), parietal operculum <strong>and</strong> insula, while the medial system<br />

includes the medial thalamus, anterior cingulate cortex, prefrontal<br />

cortex, amygdala, hippocampus, hypothalamus, nucleus accumbens,<br />

<strong>and</strong> periaqueductal grey matter (i. e. mainly components of<br />

the “limbic system” that are known to be connected to affect). In<br />

addition, there exist connections of both systems to motor-related<br />

brain regions such as primary motor cortex, striatum <strong>and</strong> cerebellum,<br />

as they are also part of the <strong>pain</strong> matrix (Peyron et al. 2000).<br />

It has been proposed that the lateral <strong>and</strong> medial <strong>pain</strong> systems<br />

have distinctive roles in producing <strong>pain</strong> perception: Activation<br />

of the lateral <strong>pain</strong> system is thought to be related to the sensorydiscriminative<br />

aspects of <strong>pain</strong>, while the medial system is believed<br />

to participate primarily in the emotional-cognitive aspects (Peyron<br />

et al. 2000, Price 2000). Already at the thalamic level, a subdivision<br />

into a sensory <strong>and</strong> emotional-cognitive part appears to be<br />

realized, as specific thalamic nuclei are interconnected with the<br />

lateral system <strong>and</strong> other nuclei with the medial system (Millan<br />

1999, Perl 2007). Those thalamic nuclei which belong to the medial<br />

system are known to play a role in affective <strong>and</strong> motivational<br />

reactions, while those of the lateral system are involved in somatosensory<br />

functions.<br />

Pain Matrix<br />

cortex<br />

spinal cord<br />

nociceptors<br />

sensorydiscriminative<br />

thalamus, brain stem,<br />

subcortex<br />

affectivecognitive<br />

Figure 4: Sensory, affective <strong>and</strong><br />

cognitive processing of nociceptive<br />

input in the brain results in <strong>pain</strong><br />

sensation<br />

The – at least partly artificial – distinction is an attempt to discriminate<br />

between those elements of <strong>pain</strong> perception which respond<br />

primarily to stimulus intensity, from those parts which reflect primarily<br />

the emotional side (“unpleasantness”) of the <strong>pain</strong> sensation.<br />

Support for the concept of an at least partial division of <strong>pain</strong> sensation<br />

into perceptions of different quality comes from clinical observations<br />

which show that selective lesions on the anterior cingulate<br />

cortex alleviate the affective component of <strong>pain</strong>, while lesions to<br />

SI/SII cortical regions affect the discriminative component of <strong>pain</strong><br />

26 <strong>Fish</strong>. <strong>Nociception</strong> <strong>and</strong> <strong>pain</strong> | Contributions to Ethics <strong>and</strong> Biotechnology<br />

<strong>Fish</strong>. <strong>Nociception</strong> <strong>and</strong> <strong>pain</strong> | Contributions to Ethics <strong>and</strong> Biotechnology<br />

27

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