1. Psychophysical experiments were initiated to determine the possible influence of increasing stimulus size on perceived pain intensity. Six trained human subjects (5 male, 1 female) made visual analogue scale (VAS) ratings for pain-sensation intensity and unpleasantness in response to nociceptive thermal stimuli. Test stimuli consisted of 5-s duration heat pulses (45-50 degrees C in 1 degrees increments) delivered by one, two, or three contact thermal probes (1 cm2 each) applied to the medial aspect of the anterior forearm. 2. The area of skin receiving noxious thermal stimuli was changed by randomly varying the number of thermodes activated. The effects of varying the distance between the thermal probes also were evaluated. In the first series of experiments, thermal-probe separation was kept close to 0; in subsequent experimental series, the thermodes were separated by either 5 or 10 cm. 3. In each experimental series, considerable spatial summation occurred in both pain-sensation intensity and unpleasantness dimensions of pain. This summation occurred throughout the nociceptive thermal range of 45-50 degrees C and was larger at suprathreshold temperatures (greater than or equal to 47 degrees C) than those near threshold (less than or equal to 46 degrees C). Unlike spatial summation of perceived warmth, that of pain was not characterized by systematic changes in power-function exponents but as approximately upward parallel displacements in double-logarithmic coordinates. 4. Thermal-probe separation over a range of 0-10 cm had no effects on spatial summation of pain-sensation intensity or pain unpleasantness. In contrast, increasing thermal-probe separation increased the subjects' ability to discriminate differences in stimulus size and their ability to detect correctly the number of thermal probes activated. 5. Because affective VAS ratings of unpleasantness were linearly related to, but distinctly and systematically less than, VAS ratings of pain-sensation intensity, it was clear that subjects responded quite differently to these two pain dimensions. Affective judgements were not additionally influenced by thermal probe separation and hence by the ability to perceive stimulus size or number of thermal probes activated. 6. The results indicate that powerful spatial-summation mechanisms exist for heat-induced pain. Spatial summation of pain is likely to be subserved both by local integration mechanisms at the level of single spinothalamic-tract neurons and by recruitment of central nociceptive neurons, because spatial summation of pain occurred to approximately equal extents under conditions of thermode separations over a distance of at least 20 cm.

A good deal of effort has been directed toward determining the sensory cues employed by neonatal animals in orienting to, and localizing, the nipple. The results of previous studies are contradictory. Some investigators have claimed that olfactory cues are critical, while others suggest that tactile cues are of primary importance in these behaviors. The present studies indicate that, in kittens, there are two essential components of the suckling process: orientation to the mother and localization of the nipple. In these experiments, the ability and time involved in localizing and attaching to the nipple of the anesthetized mother were measured in several conditions. With tactile input impaired, kittens had no difficulty locating the mother, but could not locate the nipple. In contrast, olfactory disruption never impaired nipple localization and attachment when the kitten was in contact with the mother, but interfered with the kitten's ability to locate the mother when removed from her. These data suggest that olfactory cues are employed in locating the mother, and tactile cues from the perioral region are employed in locating the nipple.

There are many somatosensory neurons in the hamster superior colliculus (SC); some respond to innocuous tactile stimuli, while others respond either preferentially, or solely, to noxious stimuli. Yet, there are little quantitative data describing the responses of these neurons. We sought to provide such information by relating stimulus intensity to the magnitude of the neural response using controlled innocuous and noxious mechanical and thermal stimuli. Of 122 somatosensory SC neurons studied in urethane-anesthetized hamsters, the majority (52%) had low-threshold mechanoreceptive properties (LT). LT neurons had force thresholds less than 1 gm, adapted rapidly to maintained stimuli, and did not respond with higher numbers of impulses to noxious mechanical or thermal stimuli. A smaller, though substantial, proportion of neurons (45%) responded either preferentially, or solely, to noxious stimuli. A few neurons (3%) were inhibited by either light tactile or noxious mechanical stimuli. Two populations of nociceptive neurons were found and classified either as wide dynamic range (WDR) neurons (n = 25), those that responded to gentle mechanical, noxious mechanical, and/or thermal stimuli; or nociceptive-specific (NS) neurons (n = 30), those that responded solely to high-intensity mechanical or noxious thermal stimuli. WDR neurons responded monotonically to increases in the intensity of innocuous mechanical stimuli, and displacement-response relationship for this population was a slightly negatively accelerating power function with an exponent of 0.785. However, the thermal stimulus-response relationships (to graded skin temperatures) of both WDR and NS neurons were positively accelerating power functions with exponents of 2.3 and 2.5 (r2 = 0.988), respectively. These values are consistent with both electrophysiological data from dorsal horn nociceptive neurons and from human psychophysical results using the same range of thermal stimuli. These experiments demonstrate that SC neurons are capable of signaling not only the presence and location of a noxious stimulus but its intensity as well. Presumably, these neurons play a significant role in the animal's reactions to potentially harmful stimuli. The partial laminar segregation of WDR and NS neurons may reflect different involvements of particular nociceptive subtypes in the various overt responses mediated by the SC.

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There is little doubt that trigeminal nociceptive neurons play a critical role in signaling the presence of harmful, or potentially harmful, orofacial stimuli. Unfortunately, there is only a limited understanding of how these neurons code such stimuli and whether this code is maintained in those structures responsible for generating overt reactions. The present series of experiments were designed to quantitatively document the response properties of nociceptive neurons in the rat trigeminal pars caudalis using the same electrical and innocuous and/or noxious mechanical and thermal stimuli employed in the characterization of nociceptive neurons with orofacial receptive fields in the rat superior colliculus. Neurons were classified as either low-threshold mechanoreceptive, wide-dynamic-range, or nociceptive-specific (type I, II) depending on their responsiveness to these stimuli. Nociceptive pars caudalis neurons (92/135, 68%) had receptive field organizations and input fibers (as indicated by latencies to electrical stimuli) quite different from those of low-threshold neurons (43/135, 32%). Nociceptive stimulus-response relationships for the population of wide-dynamic-range and nociceptive-specific type I neurons to contact heat stimuli were positively accelerating power functions with exponents of 3.9 and 4.4, respectively. This contrasted sharply with the low-threshold component of wide-dynamic-range neurons which was a negatively accelerating power function with an exponent of 0.7. All categories of nociceptive neurons also responded vigorously to cold stimuli. The thresholds of both hot and cold stimuli were often below psychophysical estimates of thermal pain, suggesting that "nociceptive" neurons process far more information than that required to signal potentially harmful stimuli. The fundamental similarities in nociceptive properties in pars caudalis and other structures of the central nervous system suggest that there is little transformation of the information encoded at successive levels of the neuraxis. This is consistent with the idea that the functional role of nociceptive neurons is reflected more in which circuits they are integrated and less in differences in their physiological properties.