Infrared thermography was used to image changes in cutaneous temperature during a conditioned fear response to context. Changes in heart rate, arterial pressure, activity and body (i.p.) temperature were recorded at the same time by radio-telemetry, in addition to freezing immobility. A marked drop in tail and paws temperature (-5.3 and -7.5 degrees C, respectively, down to room temperature), which lasted for the entire duration of the response (30 min), was observed in fear-conditioned rats. In sham-conditioned rats, the drop was on average half the magnitude and duration. In contrast, temperature of the eye, head and back increased (between + 0.8 and + 1.5 degrees C), with no difference between the two groups of rats. There was a similar increase in body temperature although it was slightly higher and delayed in the fear-conditioned animals. Finally, ending of the fear response was associated with a gradual decrease in body temperature and a rebound increase in the temperature of the tail (+ 3.3 degrees C above baseline). This study shows that fear, and to some extent arousal, evokes a strong cutaneous vasoconstriction that is restricted to the tail and paws. This regionally specific reduction in blood flow may be part of a preparatory response to a possible fight and flight to reduce blood loss in the most exposed parts of the rat's body in case of injury. The data also show that the tail is the main part of the body used for dissipating internal heat accumulated during fear once the animal has returned to a safe environment.
Hypocretin/orexin has a well-established role in wakefulness and in the maintenance of arousal. Because stress is associated with arousal, it has been proposed that hypocretin is also involved in stress. However, it is not clear if this is true for all forms of stress. To clarify this issue, we compared four conditions combining high arousal with no or low stress (wakefulness and exploration) or high stress (contextual fear and restraint) in the rat. We looked at Fos expression in hypocretin neurons, hypocretin-1 levels in cerebrospinal fluid and cardiovascular and behavioural changes after pharmacological blockade with the dual hypocretin receptor antagonist, almorexant. Fos expression in hypocretin neurons was highest with wakefulness and exploration, also high with fear but not significant with restraint. Hypocretin-1 levels were consistent with this pattern, although the differences were not as marked. Hypocretin receptor blockade with almorexant reduced the pressor, tachycardic and locomotor responses of wakefulness and exploration as well as the pressor and sympathetic component of the tachycardic response of fear. In contrast, almorexant did not reduce the pressor and tachycardic responses of restraint and nor did it reduce the pressor, tachycardic and locomotor responses of another stressor, i.e. cold exposure. Thus, hypocretin is not involved in all forms of stress. Comparison of the different conditions suggests that, regardless of stress, hypocretin involvement occurs when the arousal associated with the response includes increased attention to environmental cues. When it does, hypocretin will at least contribute to the cardiovascular response. The findings are of clinical relevance to some forms of psychological stress.
1. The contribution of the midbrain periaqueductal gray (PAG) to the central regulation of vocalization was investigated by analyzing the electromyographic (EMG) changes in respiratory, laryngeal, and oral muscles evoked by microinjection of D,L-homocysteic acid (DLH) in the PAG of unanesthetized, precollicular decerebrate cats. Moderate to large (6-40 nmol) doses of DLH evoked natural-sounding vocalization as well as increases in inspiratory depth and respiratory rate. 2. Two basic types of vocalization were evoked, each associated with a distinct and characteristic pattern of respiratory, laryngeal and oral EMG changes. Type A vocalization (voiced sounds such as howl/mew/growl) was characterized by excitation of the cricothyroid (CT) and thyro-arytenoid (TA) muscles, and inhibition of the posterior crico-arytenoid (PCA) muscle, whereas type B vocalization (unvoiced hiss sounds) was characterized by excitation of the PCA and TA muscles and no significant activation of the CT muscle. In addition, stronger expiratory (external oblique, internal oblique, internal intercostal) EMG increases were associated with type A responses, and larger increases in genioglossus and digastric muscle activity were associated with type B responses. 3. Microinjections of small doses of DLH (300 pmol-3 nmol), also evoked patterned changes in muscle activity (usually without audible vocalization) that, although of lower amplitude, were identical to those evoked by injections of moderate to large DLH doses. In no such experiments (175 sites) were individual muscles activated by small dose injections of DLH into the PAG. Further, type A vocalization/muscle patterns were evoked from PAG sites caudal to those at which type B vocalization/muscle patterns were evoked. 4. Considered together these results indicate: that the PAG contains topographically separable groups of neurons that coordinate laryngeal, respiratory, and oral muscle patterns characteristic of two fundamental types of vocalization and that the underlying PAG organization takes the form of a representation of muscle patterns, rather than individual muscles. 5. The patterns of EMG activity evoked by excitation of PAG neurons were strikingly similar to previously reported patterns of EMG activity characteristic of major phonatory categories in higher species, including humans (e.g., vowel phonation, voiceless consonant phonation). These findings raise the possibility that the sound production circuitry of the PAG could well be utilized by cortical and subcortical "language structures" to coordinate basic respiratory and laryngeal motor patterns that are necessary for speech.
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