Intensity Of Shivering Research Articles

Suppression of Shivering during Hypothermia Using a Novel Drug Combination in Healthy Volunteers

Hypothermia may be beneficial in stroke victims; however, it provokes vigorous shivering. Buspirone and dexmedetomidine each linearly reduce the shivering threshold with minimal sedation and no respiratory depression. This study tested the hypotheses that the combination of buspirone and dexmedetomidine would (1) synergistically reduce the shivering threshold, (2) synergistically reduce the gain and maximum intensity of shivering, and (3) produce sufficient inhibition to permit cooling to 34 degrees C without excessive hypotension or sedation. Eight healthy men were randomly assigned on 4 days to (1) no drug, (2) buspirone (60 mg orally), (3) dexmedetomidine (intravenous infusion to target plasma concentration of 0.6 ng/ml), or (4) combination of buspirone and dexmedetomidine at same doses. Lactated Ringer's solution (approximately 3 degrees C) was infused intravenously to decrease tympanic membrane temperature by 1.5 degrees C/h. Shivering threshold was defined as an increase in oxygen consumption greater than 20%. Sedation was evaluated using the Observer's Assessment of Sedation/Alertness scale. Mean arterial pressure and heart rate were slightly lower on dexmedetomidine and combination days. Likewise, the level of sedation was statistically different on these 2 days but clinically unimportant. Buspirone reduced the shivering threshold from 36.6 degrees C +/- 0.4 degrees C to 35.9 degrees C +/- 0.4 degrees C, dexmedetomidine reduced it to 34.7 degrees C +/- 0.5 degrees C, and the combination to 34.1 +/- 0.4 degrees C. The interaction effect of 0.04 degrees C was not significant. The gain of shivering and maximum shivering intensity were similar on each day. The combination of buspirone and dexmedetomidine additively reduced the shivering threshold. Thus, supplementing dexmedetomidine with buspirone blocks shivering and causes only minimal sedation.

Thermal responses for men with different fat compositions during immersion in cold water at two depths: prediction versus observation

A cold thermoregulatory model (CTM) was applied to data from partially immersed subjects divided into normal (NF) or low fat (LF) groups in order to validate CTM during immersion at two depths and to examine mechanisms underlying the individual differences. CTM defines thermal characteristics, e.g. surface area and maximal shivering intensity, using height, weight, fat %, age and VO(2max). Ten clothed subjects, 5 NF (15-19%) and 5 LF (8.1-14.7%), were immersed in both 10 and 15 degrees C water at chest (CH) and waist (WA) level. Environmental and clothing inputs for CTM were weighted to adjust for the ratio of skin surface area covered by either air or water at various immersion depths. Predicted core temperature (Tc) responses for each individual trial were compared with measured data. There were no significant differences (P > 0.05) between measured Tc and predicted Tc for NF at all four conditions. In contrast, for the LF group, the predicted Tc responses were all higher than measured (P < 0.05). However, predicted Tc agreed closer with measured Tc for LF when leg muscle blood flow was increased in the simulation. This suggests that blood flow may contribute to the rapid decline in Tc observed in LF and its variance may cause in part the individual differences in Tc responses. CTM predicts Tc responses to immersion at various depths with acceptable accuracy for NF individuals in this study and can be adapted to non-uniform environments.

Effect of tropisetron versus placebo on cold-induced oxygen consumption and shivering in male volunteers

In a prospective, randomized, cross-over, placebo-controlled study in healthy male volunteers, we tested the effect of the 5-HT 3 antagonist tropisetron on cold-induced oxygen consumption and shivering. Cooling was performed by intravenous infusion of isotonic salt solution at 4 °C. Whole-body oxygen consumption (VO 2) was measured with a metabolic monitor. Shivering was qualitatively assessed. When the shivering score evaluated “2” (intense shivering), 5 mg tropisetron or NaCl 0.9% was injected and repeated if necessary, to completely stop shivering. The VO 2 before shivering (178±9 ml/min/m 2) rose significantly during shivering (291±21 ml/min/m 2). 5 mg of tropisetron in 2 volunteers and 10 mg in 3 volunteers stopped shivering but it returned (grade 0–1). The VO 2 decreased significantly (209±17 ml/min/m 2). Placebo had no effect. Tropisetron reduced cold-induced VO 2 and intensity of hypothermic shivering. That an additional dose of tropisetron could not stop the shivering totally may indicate that the effect of tropisetron is not dose dependent.

Effects of carbohydrate availability on sustained shivering II. Relating muscle recruitment to fuel selection.

The purpose of this study was to quantify how shivering activity would be affected by large changes in fuel metabolism (see Haman F, Peronnet F, Kenny GP, Doucet E, Massicotte D, Lavoie C, and Weber J-M, J Appl Physiol 96: 000-000, 2004). Adult men were exposed to 10 degrees C for 2 h after a low-carbohydrate diet and exercise (Lo) and after high-carbohydrate diet without exercise (Hi). Using simultaneous metabolic and electromyographic (EMG) measurements, we quantified the effects of changes in fuel selection on the shivering activity of eight large muscles representing >90% of total shivering muscle mass. Contrary to expectation, drastic changes in fuel metabolism [carbohydrates 28 vs. 65% of total heat production (Hprod), lipids 53 vs. 23% Hprod, and proteins 19 vs. 12% Hprod for Lo and Hi, respectively] are achieved without altering the EMG signature of shivering muscles. Results show that total shivering activity and the specific contribution of each muscle to total shivering activity are not affected by large changes in fuel selection. In addition, we found that changes in burst shivering rate ( approximately 4 bursts/min), relative contribution of burst activity to total shivering ( approximately 10% of total shivering activity), and burst shivering intensity ( approximately 12% of maximal voluntary contraction) are the same between Lo and Hi. Spectral analysis of EMG signals also reveals that mean frequencies of the power spectrum remained the same under all conditions (whole body average of 78 +/- 5 Hz for Lo and 83 +/- 7 Hz for Hi). During low-intensity shivering, humans are therefore able to sustain the same thermogenic rate by oxidizing widely different fuel mixtures within the same muscle fibers.

Shivering endurance and fatigue during cold water immersion in humans.

An important component of survival time during cold exposure is shivering endurance. Nine male and three female healthy and fit subjects [mean (SD) age 24.8 (6.3) years, body mass 71.7 (13.2) kg, height 1.75 (0.10) m, body fat 22.7 (7.4)%] were immersed to the upper chest level in cold water for periods ranging from 105 to 388 min on two occasions to test a prediction of shivering endurance. The water was cooled from 20 to 8 degrees C during the first 15 min of immersion and subsequently rewarmed (<20 degrees C) to elicit a near constant submaximal shivering response. The data were divided according to moderate (M) and high (H) levels of shivering intensity. Respective mean total immersion times were 250 (75) and 199 (80) min ( P=0.086) at different average shivering intensities of 61 (10) and 69 (8)% relative to maximal shivering ( P<0.001). Blood plasma glucose concentration increased during the immersion [from 3.44 (0.54) pre- to 3.94 (0.60) mmol x l(-1) post-immersion ( P=0.037)] and levels were higher during M ( P=0.012). When compared to a model prediction of shivering endurance, shivering activity continued well beyond the predicted endurance times in 18 out of the 24 trials. The average rates of oxygen consumption over the entire immersion period were lower ( P=0.002) during M [0.93 (0.20) l x min(-1)] compared to H [1.05 (0.21) l x min(-1)), and while these rates did not change during the last 90 min of immersion, there was an increase in fat oxidation. There were no trial differences in the average esophageal (T(es)) and mean skin temperatures during the entire immersion period (36.0 and 18.0 degrees C, respectively), yet T(es) decreased ( P=0.003) approximately 0.4 degrees C during the last 90 min of immersion. When the shivering intensity was normalized to account for this decrease, a significant downward trend of approximately 17% x h(-1) in the normalized shivering intensity was found after the predicted end of shivering endurance. These results suggest that shivering drive, and not shivering intensity per se, decreased during the latter stages of the immersion. Underlying mechanisms such as fatigue and habituation for this diminishing cold sensitivity are discussed.

Measurement and prediction of peak shivering intensity in humans.

Prediction equations of shivering metabolism are critical to the development of models of thermoregulation during cold exposure. Although the intensity of maximal shivering has not yet been predicted, a peak shivering metabolic rate (Shivpeak) of five times the resting metabolic rate has been reported. A group of 15 subjects (including 4 women) [mean age 24.7 (SD 6) years, mean body mass 72.1 (SD 12) kg, mean height 1.76 (SD 0.1) m, mean body fat 22.3 (SD 7)% and mean maximal oxygen uptake (VO2max) 53.2 (SD 9) ml O2.kg-1.min-1] participated in the present study to measure and predict Shivpeak. The subjects were initially immersed in water at 8 degrees C for up to 70 min. Water temperature was then gradually increased at 0.8 degree C.min-1 to a value of 20 degrees C, which it was expected would increase shivering heat production based on the knowledge that peripheral cold receptors fire maximally at approximately this temperature. This, in combination with the relatively low core temperature at the time this water temperature was reached, was hypothesized would stimulate Shivpeak. Prior to warming the water from 8 to 20 degrees C, the oxygen consumption was 15.1 (SD 5.5) ml.kg-1.min-1 at core temperatures of approximately 35 degrees C. After the water temperature had risen to 20 degrees C, the observed Shivpeak was 22.1 (SD 4.2) ml O2.kg-1.min-1 at core and mean skin temperatures of 35.2 (SD 0.9) and 22.1 (SD 2.2) degrees C, respectively. The Shivpeak corresponded to 4.9 (SD 0.8) times the resting metabolism and 41.7 (SD 5.1)% of VO2max. The best fit equation predicting Shivpeak was Shivpeak (ml O2.kg-1.min-1) = 30.5 + 0.348 x VO2max (ml O2.kg-1.min-1) - 0.909 x body mass index (kg.m-2) - 0.233 x age (years); (P = 0.0001; r2 = 0.872).

The space blanket and shivering during extradural analgesia in labour

The incidence of shivering in women during epidural analgesia in labour is up to 33%, and may be highly distressing. Its mechanism is unclear, but the most widely held theory is that it is thermogenic. Pharmacological suppression may adversely effect the foetus or induce maternal hypothermia, and although physical remedies such as direct heat application are effective, they are also expensive and cumbersome. We recently found that the space blanket, a simple aluminised metallic foil, used pre-emptively, reduces shivering after general anaesthesia. We investigated the effect of the space blanket on the incidence and intensity of shivering, axilliary skin temperature, and subjective perception of cold during epidural analgesia for labour in 50 women over a four-hour time frame. Patients were randomised into group 1, who were wrapped in a space blanket immediately after commencement of epidural analgesia with bupivacaine, and group 2 (matched controls). Shivering was defined as visible tremor of the head, neck, trunk or limbs as observed by the attending midwives. Twenty-nine percent of group 1 and 35% of group 2 shivered (not significant), but a reduction in shivering intensity was observed in group 1 (P < 0.05). There was no significant difference in skin temperatures in either group, and no significant temperature change within the groups. Both groups had similar thermal comfort scores throughout the study, which correlated poorly with the presence of shivering. We conclude that there is no benefit in application of the space blanket to reduce epidural-analgesia related shivering in labour, which may indicate a non-thermogenic mechanism for this phenomenon.

THE ONTOGENY OF SHIVERING THERMOGENESIS IN THE RED-WINGED BLACKBIRD (AGELAIUS PHOENICEUS)

The ontogeny of shivering thermogenesis was investigated in the altricial red-winged blackbird (Agelaius phoeniceus). Two indices of heat production &shy; the rate of oxygen consumption (V(dot)O2) of the bird and the electromyographic (EMG) activity of the pectoralis (PECT) and gastrocnemius (GAST) muscles &shy; were measured simultaneously in adult and nestling red-winged blackbirds as they were subjected first to thermoneutral temperatures and subsequently to progressively colder ambient temperatures (Ta). The ontogenetic changes in both indices indicated that the capability for thermogenesis in nestling red-winged blackbirds improved markedly with age. The metabolic rates of 3-day-old nestlings decreased during exposure to gradually falling ambient temperatures; at best, these nestlings were only able to maintain mass-specific V(dot)O2 at levels similar to or slightly above the resting metabolic rate at thermoneutral temperatures (RMR) for a short time before metabolic rates decreased with further cooling. Shivering was detected only in the PECT muscles and was of a relatively low intensity (maximum of sevenfold increase in intensity over basal levels). The 5-day-old nestlings increased mass-specific V(dot)O2 modestly (approximately 1.4-fold) above RMR and attained slightly higher maximal factorial increases in the EMG activity of the PECT (maximum of 18-fold basal levels) when exposed to the same experimental conditions. Shivering was also detected in the GAST muscles of these birds. The most striking improvements in both measures observed during the nestling period occurred between day 5 and day 8. Eight-day-old nestlings increased metabolic rates by approximately 2- to 2.5-fold over basal levels and sustained these elevated rates for longer before becoming hypothermic. Both the PECT and GAST muscles contributed significantly to shivering thermogenesis, and these older nestlings attained much higher factorial increases in the intensity of shivering (up to 72-fold) during exposure to cold temperatures. In addition, both the range and magnitude of the dominant frequencies of muscle activity in the PECT increased during postnatal development. The PECT muscles were a principal site of shivering thermogenesis in all nestling and adult red-winged blackbirds studied here. Shivering in these muscles was a 'first line defense' against cold; the threshold temperature for shivering in the PECT muscles coincided with the lower critical temperature for oxygen consumption (TLC), and the subsequent increases in EMG activity in this muscle with further cooling correlated well with the corresponding increases in mass-specific V(dot)O2.

The intensity of postoperative shivering is unrelated to axillary temperature.

The relationship between axillary temperature and postoperative shivering was examined in 302 patients who entered one recovery room in the Derbyshire Royal Infirmary over a one-month period. No relationship was found between temperature and the occurrence of shivering, or between conscious level and the occurrence of shivering.

Synchrony between breathing and shivering in three muscles of bantam hens exposed to cold eggs

Unrestrained, incubating bantam hens (Gallus domesticus) were exposed to cold, water-perfused eggs to induce shivering. Mean rectified electromyogram (EMG activity) was recorded from the iliotibialis, gastrocnemius, and pectoralis muscles, together with breathing and O2 consumption. EMG activity in all three muscles varied in synchrony with breathing, as confirmed by spectral analysis. The highest shivering intensity occurred during end expiration. This synchrony was also present when respiratory frequency was increased by hypercapnic/hypoxic stimuli and tended to improve with increasing respiratory frequency and tidal volume. The absolute modulation of shivering intensity remained relatively constant with increasing average intensity, indicating that breathing-modulated motor units are activated first. In the less active pectoralis muscle, all activity was modulated. Modulation is probably not restricted to one type of muscle fiber because it occurs in both predominantly red hindquarter muscles and the white pectoralis muscle. It is tentatively suggested that respiratory drive-related influence from the brain stem has a synchronizing effect on spinal cord activity. A bursting activity pattern may be more favorable for muscle circulation than a tonic discharge pattern.

Ventilatory and metabolic responses to cold and hypoxia in intact and carotid body-denervated rats.

The effects of hypoxia on thermoregulation and ventilatory control were studied in conscious rats before and after carotid denervation (CD). Measurements of metabolic rate (VO2), ventilation (V), shivering intensity (SI), and colonic temperature (Tc) were made in groups of eight rats subjected to three protocols. In protocols 1 and 2, at ambient temperature (Ta) of 25 and 5 degrees C, respectively, rats were exposed to normoxia and hypoxia [inspired O2 fraction (FIO2) 0.13-0.11]. In protocol 3, Ta was decreased from 25 to 5 degrees C in 30-min steps of 5 degrees C. Recordings were made in normoxia and hypoxia (FIO2 0.12). The results show that in both intact and CD rats 1) in normoxia, cold exposure increased VO2, V, and SI, and these increases were proportional to the decrease in Ta; 2) hypoxia induced only a transient decrease in SI, and, for a given Ta, VO2 was reduced whereas V and SI were increased; and 3) in CD rats, V increased less during cold exposure in both normoxia and hypoxia; VO2 and Tc were more depressed during hypoxia. It is concluded that 1) the interaction between Ta and FIO2 in the control of V is partly dependent on the carotid body afferents, 2) shivering thermogenesis may be transiently affected by hypoxia independently of the carotid body afferents, and 3) nonshivering thermogenesis may be directly inhibited by hypoxia, especially during cold exposure.

Relative intensity of muscular contraction during shivering.

The intensity of cold-induced shivering, quantified by surface electromyography (EMG) and then expressed as a function of the maximal myoelectrical activity (integrated EMG) obtained during a maximum voluntary contraction (MVC), was examined in this study in individuals classified by body fat. In addition, the relationship between shivering and metabolic rate (MR) and the relative contribution of various muscle groups to total heat production were studied. Ten seminude male volunteers, 5 LEAN (less than 11% body fat) and 5 NORM (greater than 15% body fat) were exposed to 10 degrees C air for 2 h. EMG of six muscle groups (pectoralis major, rectus abdominis, rectus femoris, gastrocnemius, biceps brachii, and brachioradialis) was measured and compared with the EMG of each muscle's MVC. A whole body index of shivering, determined from the mass-weighted intensity of shivering of each muscle group, was correlated with MR. After the initial few minutes of exposure, only the pectoralis major, rectus femoris, and biceps brachii continued to increase their intensity of shivering. Shivering intensity was higher in the central muscles, ranging from 5 to 16% of MVC compared with that in the peripheral muscles, which ranged from 1 to 4% of MVC. Shivering intensities were similar in the peripheral muscles for the LEAN and NORM groups, whereas differences occurred in the trunk muscles for the pectoralis major and rectus abdominis. The whole body index of shivering correlated significantly with each individual's increase in MR (r = 0.63-0.97).(ABSTRACT TRUNCATED AT 250 WORDS)

Control of metabolic and ventilatory responses to cold in anesthetized cats

Interactions between the control of thermogenesis and ventilation were studied during normoxia, hyperoxia, and ambient or CO hypoxia in adult anesthetized intact or carotid-denervated cats. Shivering, metabolic and ventilatory responses to cold stress were studied. In addition, the effects of transient pharmacological stimulation (NaCN) or inhibition (Dopamine) of arterial chemoreceptor activity were studied under different levels of oxygenation. In intact animals, cold exposure provoked increases in V̇ O2 and ventilation which were directly proportional to the intensity of shivering. During ambient or CO hypoxia, V̇ O2 was less than in normoxia for all values of shivering intensity, suggesting that a non-shivering thermogenesis component may also be inhibited by hypoxia. The decrease in V̇ O2 was associated with a smaller decrease in ventilation in ambient than in CO hypoxia because of the presence of the chemoreflex drive during ambient hypoxia. Pharmacological changes in chemoreceptor activity induced transient and opposite changes in ventilation and shivering intensity, confirming their role in the control of thermogenesis. After carotid denervation, when the drug effects were incosistent or absent, changes in levels of oxygenation were still followed by changes in shivering activity and associated changes in V̇ O2 and ventilation. We conclude that control of thermogenesis and ventilation and their interaction may be mediated by chemoreceptors as well as by direct effects upon central, possibly diencephalic structures.

Shivering onset, metabolic response, and convective heat transfer during cold air exposure

The onset and intensity of shivering of various muscles during cold air exposure are quantified and related to increases in metabolic rate and convective heat loss. Thirteen male subjects resting in a supine position and wearing only shorts were exposed to 10 degrees C air (42% relative humidity and less than 0.4 m/s airflow) for 2 h. Measurements included surface electromyogram recordings at six muscle sites representing the trunk and limb regions of one side of the body, temperatures and heat fluxes at the same contralateral sites, and metabolic rate. The subjects were grouped according to lean (LEAN, n = 6) and average body fat (NORM, n = 7) content. While the rectal temperatures fluctuated slightly but not significantly during exposure, the skin temperature decreased greatly, more at the limb sites than at the trunk sites. Muscles of the trunk region began to shiver sooner and at a higher intensity than those of the limbs. The intensity of shivering and its increase over time of exposure were consistent with the increase in the convective heat transfer coefficient calculated from skin temperatures and heat fluxes. Both the onset of shivering and the magnitude of the increase in metabolic rate due to shivering were higher for the LEAN group than for the NORM group. A regression analysis indicates that, for a given decrease in mean skin temperature, the increase in metabolic rate due to shivering is attenuated by the square root of percent body fat. Thus the LEAN group shivered at higher intensity, resulting in higher increases in metabolic heat production and convective heat loss during cold air exposure than did the NORM group.

Extradurals and Shivering Effects of Cold and Warm Extradural Saline Injections in Volunteers

We tested the hypothesis that cooling the extradural space may provoke shivering, by giving three 80-ml extradural injections of warm (39.8±1.2°C) or cold (17±2.2°C) saline to four healthy volunteers, whilst recording central temperature and electromyographic activity from four muscles. The first injection (always cold) did not induce shivering in any of the subjects. The second and third injections, randomly cold or warm, were given after induction of shivering with cold blankets, but had no detectable effects on the intensity of shivering. This suggests that shivering in extradural anaesthesia does not result solely from cooling of the extradural space.

EXTRADURALS AND SHIVERING: EFFECTS OF COLD AND WARM EXTRADURAL SALINE INJECTIONS IN VOLUNTEERS

We tested the hypothesis that cooling the extradural space may provoke shivering, by giving three 80-ml extradural injections of warm (39.8 +/- 1.2 degrees C) or cold (17 +/- 2.2 degrees C) saline to four healthy volunteers, whilst recording central temperature and electromyographic activity from four muscles. The first injection (always cold) did not induce shivering in any of the subjects. The second and third injections, randomly cold or warm, were given after induction of shivering with cold blankets, but had no detectable effects on the intensity of shivering. This suggests that shivering in extradural anaesthesia does not result solely from cooling of the extradural space.

Effects of hypoxia on metabolic rate of conscious adult cats during cold exposure.

Oxygen consumption (VO2) and shivering movements were recorded in adult, conscious cats in a thermoneutral (24-27 degrees C) and in a cold (3-8 degrees C) environment during normoxia, hypoxia, or hyperoxia for 55 min. In the cold environment, VO2 correlated with shivering index (SI) under conditions of normoxia or ambient hypoxia (FIO2 = 0.12). During normoxia, VO2 was 63% higher in the cold than the thermoneutral environment. Ambient hypoxia acutely reduced VO2 in cold and thermoneutral environments, the decrement being greater for the former than the latter. Similarly, the variation in VO2 for unit change in SI was greater in hypoxia than normoxic conditions, suggesting that hypoxia influenced nonshivering as well as shivering components of cold-induced VO2. Hypoxia induced by CO (FICO = 0.002) also reduced VO2 and SI, a result that is consistent with previous results indicating that carotid body chemoreceptors do not mediate the suppression of shivering by ambient hypoxia. Hyperoxia increased VO2 and SI in the cold, and the effects of both hypoxia and hyperoxia in the cold were antagonized by increasing FICO2 to 0.03. The results demonstrate that hypoxia suppresses VO2 in the cold by reducing the intensity of shivering and, probably, by an action on metabolic rate that is unrelated to cold-induced calorigenesis.

Thermoregulation during cold water immersion is unimpaired by low muscle glycogen levels.

This investigation studied the importance of muscle glycogen levels for body temperature regulation during cold stress. Physiological responses of eight euglycemic males were measured while they rested in cold (18 degrees C, stirred) water on two separate occasions. The trials followed a 3-day program of diet and exercise manipulation designed to produce either high (HMG) or low (LMG) preimmersion glycogen levels in the muscles of the legs, arms, and upper torso. Preimmersion vastus lateralis muscle glycogen concentrations were lower during the LMG trial (144 +/- 14 mmol glucose/kg dry tissue) than the HMG trial (543 +/- 53 mmol glucose/kg dry tissue). There were no significant differences between the two trials in shivering as reflected by aerobic metabolic rate or in the amount of body cooling as reflected by changes in rectal temperature during the immersions. Postimmersion muscle glycogen levels remained unchanged from preimmersion levels in both trials. Small but significant increases in plasma glucose and lactate concentration occurred during both immersions. Plasma glycerol increased during immersion in the LMG trial but not in the HMG trial. Plasma free fatty acid concentration increased during both immersion trials, but the change was apparent sooner in the LMG immersion. It was concluded that thermoregulatory responses of moderately lean and fatter individuals exposed to cold stress were not impaired by a substantial reduction in the muscle glycogen levels of several major skeletal muscle groups. Furthermore, the data suggest that, depending on the intensity of shivering, other metabolic substrates are available to enable muscle glycogen to be spared.

Cardiovascular and thermoregulatory responses to intrahypothalamically injected neurotransmitters in the pigeon

1. 1.Cardiovascular and thermoregulatory responses to injections of neurotransmitters (1–10μ g of noradrenaline, NA; 10 μ g dopamine, DA; 10 μ g serotonin, 5-HT; 5 20 μ g acetylcholine, ACh) into the anterior hypothalamus were studied in the pigeon. 2. 2.All transmitters depressed mean arterial pressure (MAP). The order of potency was ACh, 5-HT, DA and NA. All monoamines (NA, DA, 5-HT) lowered also heart rate, while an intense tachycardia was seen during ACh-induced hypotension. The cardiovascular responses for NA, DA and ACh were similar at +25°C and +12°C, while 5-HT had a significantly smaller effect at +12°C. 3. 3.All transmitter reduced shivering intensity and body temperature ( T b ) at +12°C, but the degree of suppression did not correlate to changes in MAP. The order of potency for suppression of shivering was NA, DA, ACh and 5-HT. Foot temperature measurements showed that ACh induced a significant peripheral vasolidation, which enhanced the drop in T b . 4. 4.These results show that despite the strong neuronal interaction between cardivascular and thermoregulatory pathways in the avian hypothalamus, independent responses can also be elicited by exogenous transmitters. The previously described strong inhibition of shivering by intrahypothalamically injected NA is not caused by its effects on arterial pressure, but represents a more direct response of the thermoregulatory control system.

Respiratory modulation of shivering intensity in the pigeon.

Respiration and shivering were measured in unanaesthetized, cold-exposed pigeons using pneumotachography and electromyography, respectively. The instantaneous intensity of shivering in the pectoral muscle varied in phase with respiration. Power spectral analysis showed that the main frequency components of respiration and demodulated EMG coincided exactly. The intensity of shivering was highest during end-expiration and lowest at end-inspiration. This was confirmed by cross-correlation analysis of respiration and demodulated EMG. The absolute level of modulation remained constant (c. 10 microV peak-to-peak) despite changes in the general intensity of shivering. On the other hand, the relative depth of modulation was highest during incipient shivering. These facts indicate that only a part of the motor units recruited for shivering is susceptible to respiratory modulation and that this part is first recruited during incipient shivering. Inhalation of 5% CO2 did not affect the interaction between respiration and shivering although respiration frequency varied from 25 to 60 min-1. Thus, pulmonary chemoreceptors do not mediate this effect. It is suggested that the interaction between respiration and shivering occurs directly in the CNS. The question whether the interaction is adaptive for the animal or merely reflects a common evolutionary history of the underlying neural circuits is discussed.