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)     

Thermoregulatory model for immersion of humans in cold water

The mathematical models of thermoregulation of Stolwijk and Hardy, and Montgomery were used to develop a model suitable for the simulation of human physiological responses to cold-water immersion. Data were obtained from experiments where 13 healthy male volunteers were totally immersed under resting and nude conditions for 1 h in water temperatures of 20 and 28 degrees C. At these temperatures, the mean measured rectal temperature (Tre) fell by approximately 0.9 and 0.5 degrees C, respectively, yet mean measured metabolic rate (M) rose by approximately 275 and 90 W for the low body fat group (n = 7) and 195 and 45 W for the moderate body fat group (n = 6). To predict the observed Tre and M values, the present model 1) included thermal inputs for shivering from the skin independent of their inclusion with the central temperature to account for the observed initial rapid rise in M, 2) determined a thermally neutral body temperature profile such that the measured and predicted initial values of Tre and M were matched, 3) confined the initial shivering to the trunk region to avoid an overly large predicted initial rate of rectal cooling, and 4) calculated the steady-state convective heat loss by assuming a zero heat storage in the skin compartment to circumvent the acute sensitivity to the small skin-water temperature difference when using conventional methods. The last three modifications are unique to thermoregulatory modeling.     

Shivering thermogenesis in the neonatal pig

(I) Shivering intensity and metabolic rate were determined in Large White pigs aged 2, 24, 48 h and 5 d, at temperatures ranging from thermoneutrality (36°C) to cold (20°C). (2) Shivering is the main heat producing mechanism, the absence of nonshivering thermogenesis being implied by both the absence of delay between the onset of shivering (S_tt) and the increase in metabolic rate (L_ct) and by the linearity of the relationship between metabolic rate and shivering intensity in the cold. (3) For a comparable thermal demand, shivering intensity decreased with age whereas cold induced heat production remained constant, which suggests that the thermogenic efficiency of shivering is improved during the first 5 days of life.     

Thermal insulation and body temperature wearing a thermal swimsuit during water immersion

This study evaluated the effects of a thermal swimsuit on body temperatures, thermoregulatory responses and thermal insulation during 60 min water immersion at rest. Ten healthy male subjects wearing either thermal swimsuits or normal swimsuits were immersed in water (26 degrees C or 29 degrees C). Esophageal temperature, skin temperatures and oxygen consumption were measured during the experiments. Metabolic heat production was calculated from oxygen consumption. Heat loss from skin to the water was calculated from the metabolic heat production and the change in mean body temperature during water immersion. Total insulation and tissue insulation were estimated by dividing the temperature difference between the esophagus and the water or the esophagus and the skin with heat loss from the skin. Esophageal temperature with a thermal swimsuit was higher than that with a normal swimsuit at the end of immersion in both water temperature conditions (p<0.05). Oxygen consumption, metabolic heat production and heat loss from the skin were less with the thermal swimsuit than with a normal swimsuit in both water temperatures (p<0.05). Total insulation with the thermal swimsuit was higher than that with a normal swimsuit due to insulation of the suit at both water temperatures (p<0.05). Tissue insulation was similar in all four conditions, but significantly higher with the thermal swimsuit in both water temperature conditions (p<0.05), perhaps due to of the attenuation of shivering during immersion with a thermal swimsuit. A thermal swimsuit can increase total insulation and reduce heat loss from the skin. Therefore, subjects with thermal swimsuits can maintain higher body temperatures than with a normal swimsuit and reduce shivering thermo-genesis.     

Shivering modulation in humans: Effects of rapid changes in environmental temperature

During cold exposure, increase in heat production is produced via the activation of shivering thermogenesis and nonshivering thermogenesis, the former being the main contributor to compensatory heat production in non-acclimatized humans. In rats, it has been demonstrated that shivering thermogenesis is modulated solely by skin thermoreceptors but this modulation has yet to be investigated in humans. The aim of this study was to determine if cold-induced shivering in humans can be modulated by cutaneous thermoreceptors in conditions where increases in heat loss can be adequately compensated by increases in thermogenic rate. Using a liquid-conditioned suit, six non-acclimatized men were exposed to cold (6 °C) for four 30 min periods, each of them separated by 15 min of heat exposure (33 °C). Core temperature remained stable throughout exposures whereas skin temperatures significantly decreased by 12% in average during the sequential cold/heat exposures compared to baseline (p<0.0001). Shivering intensity and metabolic rate increased significantly during 6 °C exposures (3.3±0.7% MVC, 0.40±0.0 L O₂/min, respectively) and were significantly reduced during 33 °C exposure (0.5±0.1% MVC, 0.25±0.0 L O₂/min; p<0.005 for both). Most importantly, shivering could be quickly and strongly inhibited during 33 °C exposure although skin temperature often remained below baseline values. In conclusion, under compensatory conditions, cutaneous thermoreceptors appear to be a major modulator of the shivering response in humans and seem to react rapidly to changes in the microclimate right next to the skin and to skin temperature.     

Effect of uniform and non-uniform skin temperature on thermal exchanges in water in humans

We investigated the effect of uniform (UST) and non-uniform (NUST) skin temperature on thermal exchanges during a 3-h water immersion in five male subjects wearing (NUST) or not wearing (UST) a water-perfused garment. UST was achieved by immersing the nude subject in water up to the neck. For each subject, the water temperature was adjusted to the critical temperature ( T(cw), 31.4 +/- 0.9 degrees C) or 3 degrees C below T(cw) ( T(cw) - 3). NUST was achieved by perfusing different segments of the perfused garment with water of different temperatures. The water temperature of the segment was independently adjusted according to the skin temperature distribution in cold air, the mean skin temperature being the same as the UST. At T(cw) and T(cw) - 3, changes in esophageal and mean skin temperatures were identical in UST and NUST conditions, but the skin temperature of the trunk was higher and that of the limb was lower in the NUST condition. Heat production and the overall skin heat flux at T(cw) were identical in the two conditions, but those at T(cw) - 3 were about 25% lower ( P < 0.05) in NUST than in UST conditions. At T(cw) - 3, the overall tissue insulation was 36% higher ( P < 0.05) in NUST than in UST conditions, mainly because of higher limb insulation. Thermogenesis due to shivering was lower by 62% ( P < 0.05) in NUST than in UST. We conclude that the NUST condition increased tissue insulation and suppressed shivering. This suggests that a high skin temperature of the trunk attenuates shivering in cold water and increases the ability to defend body temperature more economically in cold water.     

Physical fitness and thermoregulatory reactions in a cold environment in men

The relationship between the physical fitness level (maximal O2 consumption, VO2max) and thermoregulatory reactions was studied in 17 adult males submitted to an acute cold exposure. Standard cold tests were performed in nude subjects, lying for 2 h in a climatic chamber at three ambient air temperatures (10, 5, and 1 degrees C). The level of physical fitness conditioned the intensity of thermoregulatory reactions to cold. For all subjects, there was a direct relationship between physical fitness and 1) metabolic heat production, 2) level of mean skin temperature (Tsk), 3) level of skin conductance, and 4) level of Tsk at the onset of shivering. The predominance of thermogenic or insulative reactions depended on the intensity of the cold stress: insulative reactions were preferential at 10 degrees C, or even at 5 degrees C, whereas colder ambient temperature (1 degree C) triggered metabolic heat production abilities, which were closely related to the subject's physical fitness level. Fit subjects have more efficient thermoregulatory abilities against cold stress than unfit subjects, certainly because of an improved sensitivity of the thermoregulatory system.     

Human thermoregulatory responses during prolonged walking in water at 25, 30 and 35°C

Eight healthy and physically well-trained male students exercised on a treadmill for 60 min while being immersed in water to the middle of the chest in a laboratory flowmill. The water velocity was adjusted so that the intensity of exercise correspond to 50% maximal oxygen uptake of each subject, and experiments were performed once at each of three water temperatures: 25, 30, 35°C, following a 30-min control period in air at 25°C, and on a treadmill in air at an ambient temperature of 25°C. Thermal states during rest and exercise were determined by measuring rectal and skin temperatures at various points, and mean skin temperatures were calculated. The intensity of exercise was monitored by measuring oxygen consumption, and heart rate was monitored as an indicator for cardiovascular function. At each water temperature, identical oxygen consumption levels were attained during exercise, indicating that no extra heat was produced by shivering at the lowest water temperature. The slight rise in rectal temperature during exercise was not influenced by the water temperature. The temperatures of skin exposed to air rose slightly during exercise at 25°C and 30°C water temperature and markedly at 35°C. The loss of body mass increased with water temperature indicating that both skin blood flow and sweating during exercise increased with the rise in water temperature. The rise in body temperature provided the thermoregulatory drive for the loss of the heat generated during exercise. Heart rate increased most during exercise in water at 35°C, most likely due to enhanced requirements for skin blood flow. Although such requirements were certainly smallest at 25°C water temperature, heart rate at this temperature was slightly higher than at 30°C suggesting reflex activation of sympathetic control by cold signals from the skin. There was a significantly greater increase in mean skin and rectal temperatures in subjects exercising on the treadmill in air, compared to those exercising in water at 25°C. Accepted: 22 May 1998     

Temperatures of skin, subcutaneous tissue, muscle and core in resting men in cold, comfortable and hot conditions

To examine the core-shell model of temperature distribution and the possible role of subcutaneous temperature in heat regulation, comprehensive temperature measurements were made on six nude resting men exposed for 2-3 h to comfort (27 degrees C), cold (15 degrees C) and heat (45 degrees C). Cold produced strong shivering and heat caused heavy sweating. Temperatures were recorded every 10 min from: esophagus, rectum and auditory canal; back muscle and thigh muscle at 20 mm and 40 mm depths; 6 subcutaneous sites; and 16 skin sites. Average temperatures at these 29 sites were tabulated at the ends of comfort, hot and cold and the onsets of sweating and shivering. Body temperature changes were slow to develop, the skin temperatures being fastest, and successively deeper tissues progressively slower. There was occasional after-drop and after-rise. The data were consistent with the core-shell concept. The temperature gradient from subcutaneous tissue to skin, which differed substantially with comfort, the onset of shivering and the onset of sweating, could serve as a regulatory signal. The data are now in computer format and may be of interest to biothermal modelers.     

Seasonal changes in thermal responses of urban residents to cold exposure

To determine whether urban circumpolar residents show seasonal acclimatisation to cold, thermoregulatory responses and thermal perception during cold exposure were examined in young men during January-March (n=7) and August-September (n=8). Subjects were exposed for 24 h to 22 and to 10 degrees C. Rectal (T(rect)) and skin temperatures were measured throughout the exposure. Oxygen consumption (VO(2)), finger skin blood flow (Q(f)), shivering and cold (CDT) and warm detection thresholds (WDT) were assessed four times during the exposure. Ratings of thermal sensations, comfort and tolerance were recorded using subjective judgement scales at 1-h intervals. During winter, subjects had a significantly higher mean skin temperature at both 22 and 10 degrees C compared with summer. However, skin temperatures decreased more at 10 degrees C in winter and remained higher only in the trunk. Finger skin temperature was higher at 22 degrees C, but lower at 10 degrees C in the winter suggesting an enhanced cold-induced vasoconstriction. Similarly, Q(f) decreased more in winter. The cold detection threshold of the hand was shifted to a lower level in the cold, and more substantially in the winter, which was related to lower skin temperatures in winter. Thermal sensations showed only slight seasonal variation. The observed seasonal differences in thermal responses suggest increased preservation of heat especially in the peripheral areas in winter. Blunted vasomotor and skin temperature responses, which are typical for habituation to cold, were not observed in winter. Instead, the responses in winter resemble aggravated reactions of non-cold acclimatised subjects.     

Prediction of the thermoregulatory response for clothed immersion in cold water

A multi-compartmental thermoregulatory model was applied to data of ten resting clothed males immersed for 3 h in water at 10 and 15 degrees C. Clothing consisted of a dry suit and either a light or heavy undergarment, representing a total insulation of 0.15 (0.95) or 0.20 m2 degrees CW-1 (1.28 clo), respectively. Data were grouped according to low (less than 14%) and high (14 to 24%) body fat individuals. Mean decreases in rectal temperature ranged from 0.79 to 1.38 degrees C, mean decreases in the mean weighted skin temperature ranged from 6.3 to 10.2 degrees C, and mean increases in the metabolic rate ranged from 33.9 to 80.8 W. The model consists of eight segments, each representing a specific region of the body. Each segment is comprised of compartments representing the core, muscle, fat, skin, and clothing. Each compartment is assigned thermophysical values of heat conduction and heat capacitance, and with the exception of clothing, physiological values of blood flow and metabolic heat production. During cold exposure, responses are directed towards increased heat production in the form of shivering and heat conservation in the form of vasoconstriction and convective heat exchange at the vascular level. Agreement between the model predictions and the experimental observations was obtained by adjusting the parameters governing these responses.(ABSTRACT TRUNCATED AT 250 WORDS)     

Influence of menstrual cycle on shivering, skin blood flow, and sweating responses measured at night

In 10 women, external cold and heat exposures were performed both in the middle of luteal phase (L) and in the early follicular phase (F) of the menstrual cycle. Serum progesterone concentrations in L and F averaged 46.0 and 0.9 nmol X l-1, respectively. The experiments took place between 3:00 and 4:30 A.M., when the L-F core temperature difference is maximal. At neutral ambient temperature, esophageal (Tes), tympanic (Tty), rectal (Tre), and mean skin (Tsk) temperatures averaged 0.59 degrees C higher in L than in F. The thresholds for shivering, chest sweating, and cutaneous vasodilation (heat clearance technique) at the thumb and forearm were increased in L by an average of 0.47 degrees C, related to mean body temperature [Tb(es) = 0.87Tes + 0.13 Tsk] and to Tes, Tty, Tre, or Tsk. The above-threshold chest sweat rate and cutaneous heat clearances at the thumb and forearm were also enhanced in L, when related to Tb(es) or time. The metabolic rate, arm blood flow, and heart rate at thermoneutral conditions were increased in L by 5.0%, 1.1 ml X 100 ml-1 X min-1, and 4.6 beats X min-1, respectively. The concomitant increase in threshold temperatures for all autonomic thermoregulatory responses in L supports the concept of a resetting of the set point underlying the basal body temperature elevation in L. The effects of the increased threshold temperatures are counteracted by enhanced heat loss responses.     

Body temperature regulation in the rat

In loosely-restrained adult conscious rats exposed to stepwise changes in ambient temperature (T(a)) from 25 to 5 degrees C or from 20 to 35 degrees C, we have recorded body and tail temperatures, metabolic rate (VO(2)), shivering and ventilation (V). It was found that VO(2) and V vary with T(a) and show a nadir for a T(a) of 30 degrees C whereas shivering starts at 20 degrees C and increases progressively with cold exposure. T(tail) follows changes in T(a) whereas T(body) decreases slightly in cold and increases markedly in warm exposure. These results suggest that the control of T(body) interacts with the control of breathing in order to increase VO(2) during cold exposure and to facilitate evaporative respiratory heat dissipation during warm exposure.     

Partial-body exposure of human volunteers to 2450 MHz pulsed or CW fields provokes similar thermoregulatory responses

Many reports describe data showing that continuous wave (CW) and pulsed (PW) radiofrequency (RF) fields, at the same frequency and average power density (PD), yield similar response changes in the exposed organism. During whole-body exposure of squirrel monkeys at 2450 MHz CW and PW fields, heat production and heat loss responses were nearly identical. To explore this question in humans, we exposed two different groups of volunteers to 2450 MHz CW (two females, five males) and PW (65 micros pulse width, 10(4) pps; three females, three males) RF fields. We measured thermophysiological responses of heat production and heat loss (esophageal and six skin temperatures, metabolic heat production, local skin blood flow, and local sweat rate) under a standardized protocol (30 min baseline, 45 min RF or sham exposure, 10 min baseline), conducted in three ambient temperatures (T(a) = 24, 28, and 31 degrees C). At each T(a), average PDs studied were 0, 27, and 35 mW/cm2 (Specific absorption rate (SAR) = 0, 5.94, and 7.7 W/kg). Mean data for each group showed minimal changes in core temperature and metabolic heat production for all test conditions and no reliable differences between CW and PW exposure. Local skin temperatures showed similar trends for CW and PW exposure that were PD-dependent; only the skin temperature of the upper back (facing the antenna) showed a reliably greater increase (P =.005) during PW exposure than during CW exposure. Local sweat rate and skin blood flow were both T(a)- and PD-dependent and showed greater variability than other measures between CW and PW exposures; this variability was attributable primarily to the characteristics of the two subject groups. With one noted exception, no clear evidence for a differential response to CW and PW fields was found.     

A canine thermal model for simulating temperature responses of military working dogs

Military working dogs (MWDs) are often required to operate in dangerous or extreme environments, to include hot and humid climate conditions. These scenarios can put MWD at significant risk of heat injury. To address this concern, a two-compartment (core, skin) rational thermophysiological model was developed to predict the temperature of a MWD during rest, exercise, and recovery. The Canine Thermal Model (CTM) uses inputs of MWD mass and length to determine a basal metabolic rate and body surface area. These calculations are used along with time series inputs of environmental conditions (air temperature, relative humidity, solar radiation and wind velocity) and level of metabolic intensity (MET) to predict MWD thermoregulatory responses. Default initial values of core and skin temperatures are set at neutral values representative of an average MWD; however, these can be adjusted to match known or expected individual temperatures. The rational principles of the CTM describe the heat exchange from the metabolic energy of the core compartment to the skin compartment by passive conduction as well as the application of an active control for skin blood flow and to tongue and lingual tissues. The CTM also mathematically describes heat loss directly to the environment via respiration, including panting. Thermal insulation properties of MWD fur are also used to influence heat loss from skin and gain from the environment. This paper describes the CTM in detail, outlining the equations used to calculate avenues of heat transfer (convective, conductive, radiative and evaporative), overall heat storage, and predicted responses of the MWD. Additionally, this paper outlines examples of how the CTM can be used to predict recovery from exertional heat strain, plan work/rest cycles, and estimate work duration to avoid overheating.     

Repeated exposures to cold and the relationship between skin and core temperatures in control of metabolic rate in the goat (Capra hircus)

1. After 10-12 experiments in each of three goats, in which skin or core temperatures were lowered while the other temperatures remained sufficiently high to prevent metabolic rate from increasing, the core temperature threshold of shivering was lowered by 0.35 degrees C. 2. After 10-15 experiments, in which skin and core temperatures were simultaneously lowered to induce major increases of metabolic rate, no further change of threshold was observed, while the slope of metabolic rate over core temperature was reduced. 3. It is concluded that repeated cold exposures without manifest shivering can induce tolerance adaptation to cold.     

Temperature-induced changes in metabolism and body weight of cattle (Bos taurus).

The metabolic and body weight changes in two non-pregnant beef cows were studied during prolonged exposure to warm (20 +/- 3 degrees C, relative humidity 50-70%) and cold (-10 +/- 2 or -25 +/- 4 degrees C) temperatures. Other factors including daily food intake were held constant throughout each 8-week exposure. During cold exposures, metabolic rate, blood hematocrit, and plasma concentrations of glucose and free fatty acid were elevated and respiratory frequencies and skin temperatures decreased. Resting metabolic rates measured at 20 degrees C, i.e., without the direct influence of cold, were 83.4-95.3 litres 02 per hour when the cows were cold acclimated, at either -10 or -25 degrees C, and 30-40% greater than when the cows were warm acclimated. The resting metabolic response and the concomitant reduction in intensity of shivering is indicative of metabolic acclimation to cold in these animals of greater than 500 kg body weight. As well as the expected changes in body weight with changes in energy metabolism there were losses in weight (13-24 kg) during the first 3 days of each cold exposure. Weight gains occurred when the cold stress was abruptly removed. These short term weight changes were associated with changes in water intake and apparent shifts in body fluid content.     

Mechanism of action of physical antipyresis in the rat

To study the mechanism of action of physical antipyresis, core temperature was measured in two groups of rats in which heat loss was increased by cold exposure and by cooling an inferior cava heat exchanger, respectively, both before and after infection with Salmonella enteritidis. Cold exposure did not influence core temperature. On the other hand, cooling the heat exchanger caused a fall in core temperature of approximately 0.7 degree C, to 37 degrees C in normothermia and to 38.5 degrees C 24 h after the infection. These lower core temperatures were then regulated against any further increase in heat loss until the thermoregulatory metabolic capacity of the animals was exhausted and a hypothermia developed. It is concluded that in infectious fever the threshold temperature of shivering increases as much as core temperature. Furthermore it is suggested that physical antipyresis, such as sponging with tepid water, induces a moderate but regulated fall in temperature to about the threshold of shivering and that its efficacy may increase with ambient temperature.     

Involvement of basal metabolic rate in determination of type of cold tolerance

This study aimed to assess the relationship between basal metabolic rate (BMR) and metabolic heat production, and to clarify the involvement of BMR in determining the phenotype of cold tolerance. Measurements of BMR, maximum oxygen uptake, and cold exposure test were conducted on ten males. In the cold exposure test, rectal (T(rec)) and mean skin temperatures (T(ms)), oxygen uptake, and blood flow at forearm (BF(arm)) were measured during exposure to cold (10 degrees C) for 90 min. Significant correlations were observed between BMR and increasing rate of oxygen uptake, as well as between decreasing rate of BF(arm) and increasing rate of oxygen uptake at the end of cold exposure. These findings suggested that individuals with a lower BMR were required to increase their metabolic heat production during cold exposure, and that those with a higher BMR were able to moderate increased metabolic heat production during cold exposure because they were able to reduce heat loss. This study showed that BMR is an important factor in determining the phenotype of cold tolerance, and that individuals with a low BMR showed calorigenic-type cold adaptation, whereas subjects with a high BMR exhibited adiabatic-type cold adaptation by peripheral vasoconstriction.     

Effect of ambient temperature on heat production and heat loss in burn patients.

Four controls and eight burned patients with thermal injury ranging from 7 to 84% total body surface were studied in an environmental chamber at 25 and 33 degrees C ambient temperature and a constant vapor pressure during two consecutive 24-h periods. Hypermetabolism was present in the burn patients in both ambient temperatures and core and skin temperatures were consistently higher than in the normal men despite increased evaporative water loss. The higher environmental temperature decreased metabolic rate in patients with large thermal injuries in whom the decrement in dry heat loss produced by higher ambient temperature exceeded the increase of wet heat loss. In patients with burns smaller than 60%, these changes equaled one another and higher environmental temperature exerted no effect on metabolic rate. Core-skin heat conductivity increased with burn size; patients with large burns were characterized by inadequate core-skin insulation when exposed to the cooler environment, necessitating the compensatory increase of metabolic rate. This increase, however, was small and of the order of 5-8 kcal times m-2 times h-1.