Shivering Heat Production Research Articles

A geometrically accurate 3 dimensional model of human thermoregulation for transient cold and hot environments

This paper outlines the development of a finite element human thermoregulatory model using an anatomically and geometrically correct human body model. The finite element body model was constructed from digital Phantoms and is anatomically realistic, including 13 organs and tissues: skin, muscles, fat, bones, heart, lungs, brain, bladder, intestines, stomach, kidneys, liver, and eyes. The model simulates thermal responses through a passive and active system. The passive system describes heat balance within the body and between the skin surface and environment. The active system describes thermoregulatory mechanisms, i.e., vasodilation, vasoconstriction, sweating, and shivering heat production. This model predicts temperature distribution across the body at high spatial resolution, and provides insight into human thermoregulatory responses to non-uniform and transient environments. Predicted temperatures (i.e., core, skin, muscle and fat) at 29 sites were compared with measured values in comfort, hot, and cold conditions. The comprehensive validation shows predictions are accurate and acceptable.

Comparison of Electric Resistive Heating Pads and Forced-Air Warming for Pre-hospital Warming of Non-shivering Hypothermic Subjects.

Victims of severe hypothermia require external rewarming, as self-rewarming through shivering heat production is either minimal or absent. The US Military commonly uses forced-air warming in field hospitals, but these systems require significant power (600-800 W) and are not portable. This study compared the rewarming effectiveness of an electric resistive heating pad system (requiring 80 W) to forced-air rewarming on cold subjects in whom shivering was pharmacologically inhibited. Shivering was inhibited by intravenous meperidine (1.5 mg/kg), administered during the last 10 min of cold-water immersion. Subjects then exited from the cold water, were dried and lay on a rescue bag for 120 min in one of the following conditions: spontaneous rewarming only (rescue bag closed); electric resistive heating pads (EHP) wrapped from the anterior to posterior torso (rescue bag closed); or, forced-air warming (FAW) over the anterior surface of the body (rescue bag left open and cotton blanket draped over warming blanket). Supplemental meperidine (to a maximum cumulative dose of 3.3 mg/kg) was administered as required during rewarming to suppress shivering. Six healthy subjects (3 m, 3 f) were cooled on three different occasions, each in 8°C water to an average nadir core temperature of 34.4 ± 0.6°C (including afterdrop). There were no significant differences between core rewarming rates (spontaneous; 0.6 ± 0.3, FAW; 0.7 ± 0.2, RHP; 0.6 ± 0.2°C/h) or post-cooling afterdrop (spontaneous; 1.9 ± 0.4, FAW; 1.9 ± 0.3, RHP; 1.6 ± 0.2°C) in any of the 3 conditions. There were also no significant differences between metabolic heat production (S; 74 ± 20, FAW; 66 ± 12, RHP; 63 ± 9 W). Total heat gain was greater with FAW (36 W gain) than EHP (13 W gain) and spontaneous (13 W loss) warming (p < 0.005). Total heat gain was greater in FAW than both EHP, and spontaneous rewarming conditions, however, there were no observed differences found in rewarming rates, post-cooling afterdrop or metabolic heat production. The electric heat pad system provided similar rewarming performance to a forced-air warming system commonly used in US military field hospitals for hypothermic patients. A battery-powered version of this system would not only relieve pressure on the field hospital power supply but could also potentially allow extending use to locations closer to the field of operations and during transport. Such a system could be studied in larger groups in prospective trials on colder patients.

Human Responses to 5 Heated Hypothermia Wrap Systems in a Cold Environment

We compared the effectiveness of 5 heated hypothermia wrap systems. Physiologic and subjective responses were determined in 5 normothermic subjects (1 female) for 5 heated hypothermia wraps (with vapor barrier and chemical heat sources) during 60 min of exposure to a temperature of -22°C. The 5 systems were 1) user-assembled; 2) Doctor Down Rescue Wrap; 3) hypothermia prevention and management kit (HPMK); 4) MARSARS Hypothermia Stabilizer Bag; and 5) Wiggy's Victims Casualty Hypothermia Bag. Core and skin temperature, metabolic heat production, skin heat loss, and body net heat gain were determined. Subjective responses were also evaluated for whole body cold discomfort, overall shivering rating, overall temperature rating, and preferential ranking. The Doctor Down and user-assembled systems were generally more effective, with higher skin temperatures and lower metabolic heat production; they allowed less heat loss, resulting in higher net heat gain (P<0.05). HPMK had the lowest skin temperature and highest shivering heat production and scored worse than the other 4 systems for the "whole body cold discomfort" and "overall temperature" ratings (P<0.05). The user-assembled and Doctor Down systems were most effective, and subjects were coldest with the HPMK system. However, it is likely that any of the tested systems would be viable options for wilderness responders, and the choice would depend on considerations of cost; volume, as it relates to available space; and weight, as it relates to ability to carry or transport the system to the patient.

Large muscles are beneficial but not required for improving thermogenic capacity in small birds

It is generally assumed that small birds improve their shivering heat production capacity by developing the size of their pectoralis muscles. However, some studies have reported an enhancement of thermogenic capacity in the absence of muscle mass variation between seasons or thermal treatments. We tested the hypothesis that an increase in muscle mass is not a prerequisite for improving avian thermogenic capacity. We measured basal (BMR) and summit (Msum) metabolic rates of black capped chickadees (Poecile atricapillus) acclimated to thermoneutral (27 °C) and cold (−10 °C) temperatures and obtained body composition data from dissections. Cold acclimated birds consumed 44% more food, and had 5% and 20% higher BMR and Msum, respectively, compared to individuals kept at thermoneutrality. However, lean dry pectoralis and total muscle mass did not differ between treatments, confirming that the improvement of thermogenic capacity did not require an increase in skeletal muscle mass. Nevertheless, within temperature treatments, Msum was positively correlated with the mass of all measured muscles, including the pectoralis. Therefore, for a given acclimation temperature individuals with large muscles do benefit from muscle size in term of heat production but improving thermogenic capacity during cold acclimation likely requires an upregulation of cell functions.

Uncoupling Basal and Summit Metabolic Rates in White-Throated Sparrows: Digestive Demand Drives Maintenance Costs, but Changes in Muscle Mass Are Not Needed to Improve Thermogenic Capacity.

Avian basal metabolic rate (BMR) and summit metabolic rate (Msum) vary in parallel during cold acclimation and acclimatization, which implies a functional link between these variables. However, evidence suggests that these parameters may reflect different physiological systems acting independently. We tested this hypothesis in white-throated sparrows (Zonotrichia albicollis) acclimated to two temperatures (-8° and 28°C) and two diets (0% and 30% cellulose). We expected to find an uncoupling of Msum and BMR where Msum, a measure of maximal shivering heat production, would reflect muscle and heart mass variation and would respond only to temperature, while BMR would reflect changes in digestive and excretory organs in response to daily food intake, responding to both temperature and diet. We found that the gizzard, liver, kidneys, and intestines responded to treatments through a positive relationship with food intake. BMR was 15% higher in cold-acclimated birds and, as expected, varied with food intake and the mass of digestive and excretory organs. In contrast, although Msum was 19% higher in cold-acclimated birds, only heart mass responded to temperature (+18% in the cold). Pectoral muscles did not change in mass with temperature but were 8.2% lighter on the cellulose diet. Nevertheless, Msum varied positively with the mass of heart and skeletal muscles but only in cold-acclimated birds. Our results therefore suggest that an upregulation of muscle metabolic intensity is required for cold acclimation. This study increases support for the hypothesis that BMR and Msum reflect different physiological systems responding in parallel to constraints associated with cold environments.

Comparison of Distal Limb Warming With Fluidotherapy and Warm Water Immersion for Mild Hypothermia Rewarming

The purpose of the study was to determine the effectiveness of Fluidotherapy rewarming through the distal extremities for mildly hypothermic, vigorously shivering subjects. Fluidotherapy is a dry heat modality in which cellulose particles are suspended by warm air circulation. Seven subjects (2 female) were cooled on 3 occasions in 8˚C water for 60 minutes, or to a core temperature of 35°C. They were then dried and rewarmed in a seated position by 1) shivering only; 2) Fluidotherapy applied to the distal extremities (46 ± 1°C, mean ± SD); or 3) water immersion of the distal extremities (44 ± 1°C). The order of rewarming followed a balanced design. Esophageal temperature, skin temperature, heart rate, oxygen consumption, and heat flux were measured. The warm water produced the highest rewarming rate, 6.1°C·h(-1), 95% CI: 5.3-6.9, compared with Fluidotherapy, 2.2°C·h(-1), 95% CI: 1.4-3.0, and shivering only, 2.0°C·h(-1), 95% CI: 1.2-2.8. The Fluidotherapy and warm water conditions increased skin temperature and inhibited shivering heat production, thus reducing metabolic heat production (166 ± 42 W and 181 ± 45 W, respectively), compared with shivering only (322 ± 142 W). Warm water provided a significantly higher net heat gain (398.0 ± 52 W) than shivering only (288.4 ± 115 W). Fluidotherapy was not as effective as warm water for rewarming mildly hypothermic subjects. Although Fluidotherapy is more portable and technically simpler, it provides a lower rate of rewarming that is similar to shivering only. It does help decrease shivering heat production, lowering energy expenditure and cardiac work, and could be considered in a hospital setting, if convenient.

Comparison of Heat Donation Through the Head or Torso on Mild Hypothermia Rewarming

The purpose of the study was to compare the effectiveness of head vs torso warming in rewarming mildly hypothermic, vigorously shivering subjects using a similar source of heat donation. Six subjects (1 female) were cooled on 3 occasions in 8 ºC water for 60 minutes or to a core temperature of 35 ºC. They were then dried, insulated, and rewarmed by 1) shivering only; 2) charcoal heater applied to the head; or 3) charcoal heater applied to the torso. The order of rewarming methods followed a balanced design. Esophageal temperature, skin temperature, heart rate, oxygen consumption, and heat flux were measured. There were no significant differences in rewarming rate among the 3 conditions. Torso warming increased skin temperature and inhibited shivering heat production, thus providing similar net heat gain (268 ± 66 W) as did shivering only (355 ± 105 W). Head warming did not inhibit average shivering heat production (290 ± 72 W); it thus provided a greater net heat gain during 35 to 60 minutes of rewarming than did shivering only. Head warming is as effective as torso warming for rewarming mildly hypothermic victims. Head warming may be the preferred method of rewarming in the field management of hypothermic patients if: 1) extreme conditions in which removal of the insulation and exposure of the torso to the cold is contraindicated; 2) excessive movement is contraindicated (eg, potential spinal injury or severe hypothermia that has a risk of ventricular fibrillation); or 3) if emergency personnel are working on the torso.

Shivering heat production and body fat protect the core from cooling during body immersion, but not during head submersion: A structural equation model

Previous studies showed that core cooling rates are similar when only the head or only the body is cooled. Structural equation modeling was used on data from two cold water studies involving body-only, or whole body (including head) cooling. Exposure of both the body and head increased core cooling, while only body cooling elicited shivering. Body fat attenuates shivering and core cooling. It is postulated that this protection occurs mainly during body cooling where fat acts as insulation against cold. This explains why head cooling increases surface heat loss with only 11% while increasing core cooling by 39%.

Shivering Heat Production and Core Cooling During Head-In and Head-Out Immersion in 17°C Water

Many cold-water scenarios cause the head to be partially or fully immersed (e.g., ship wreck survival, scuba diving, cold-water adventure swim racing, cold-water drowning, etc.). However, the specific effects of head cold exposure are minimally understood. This study isolated the effect of whole-head submersion in cold water on surface heat loss and body core cooling when the protective shivering mechanism was intact. Eight healthy men were studied in 17 degrees C water under four conditions: the body was either insulated or exposed, with the head either out of the water or completely submersed under the water within each insulated/exposed subcondition. Submersion of the head (7% of the body surface area) in the body-exposed condition increased total heat loss by 11% (P < 0.05). After 45 min, head-submersion increased core cooling by 343% in the body-insulated subcondition (head-out: 0.13 +/- 0.2 degree C, head-in: 0.47 +/- 0.3 degree C; P < 0.05) and by 56% in the body-exposed subcondition (head-out: 0.40 +/- 0.3 degree C and head-in: 0.73 +/- 0.6 degree C; P < 0.05). In both body-exposed and body-insulated subconditions, head submersion increased the rate of core cooling disproportionally more than the relative increase in total heat loss. This exaggerated core-cooling effect is consistent with a head cooling induced reduction of the thermal core, which could be stimulated by cooling of thermosensitive and/or trigeminal receptors in the scalp, neck, and face. These cooling effects of head submersion are not prevented by shivering heat production.

Thermal effects of whole head submersion in cold water on nonshivering humans

This study isolated the effect of whole head submersion in cold water, on surface heat loss and body core cooling, when the confounding effect of shivering heat production was pharmacologically eliminated. Eight healthy male subjects were studied in 17 degrees C water under four conditions: the body was either insulated or uninsulated, with the head either above the water or completely submersed in each body-insulation subcondition. Shivering was abolished with buspirone (30 mg) and meperidine (2.5 mg/kg), and subjects breathed compressed air throughout all trials. Over the first 30 min of immersion, exposure of the head increased core cooling both in the body-insulated conditions (head out: 0.47 +/- 0.2 degrees C, head in: 0.77 +/- 0.2 degrees C; P < 0.05) and the body-exposed conditions (head out: 0.84 +/- 0.2 degrees C and head in: 1.17 +/- 0.5 degrees C; P < 0.02). Submersion of the head (7% of the body surface area) in the body-exposed conditions increased total heat loss by only 10%. In both body-exposed and body-insulated conditions, head submersion increased core cooling rate much more (average of 42%) than it increased total heat loss. This may be explained by a redistribution of blood flow in response to stimulation of thermosensitive and/or trigeminal receptors in the scalp, neck and face, where a given amount of heat loss would have a greater cooling effect on a smaller perfused body mass. In 17 degrees C water, the head does not contribute relatively more than the rest of the body to surface heat loss; however, a cold-induced reduction of perfused body mass may allow this small increase in heat loss to cause a relatively larger cooling of the body core.

Thermal effects of dorsal head immersion in cold water on nonshivering humans

Personal floatation devices maintain either a semirecumbent flotation posture with the head and upper chest out of the water or a horizontal flotation posture with the dorsal head and whole body immersed. The contribution of dorsal head and upper chest immersion to core cooling in cold water was isolated when the confounding effect of shivering heat production was inhibited with meperidine (Demerol, 2.5 mg/kg). Six male volunteers were immersed four times for up to 60 min, or until esophageal temperature = 34 degrees C. An insulated hoodless dry suit or two different personal floatation devices were used to create four conditions: 1) body insulated, head out; 2) body insulated, dorsal head immersed; 3) body exposed, head (and upper chest) out; and 4) body exposed, dorsal head (and upper chest) immersed. When the body was insulated, dorsal head immersion did not affect core cooling rate (1.1 degrees C/h) compared with head-out conditions (0.7 degrees C/h). When the body was exposed, however, the rate of core cooling increased by 40% from 3.6 degrees C/h with the head out to 5.0 degrees C/h with the dorsal head and upper chest immersed (P < 0.01). Heat loss from the dorsal head and upper chest was approximately proportional to the extra surface area that was immersed (approximately 10%). The exaggerated core cooling during dorsal head immersion (40% increase) may result from the extra heat loss affecting a smaller thermal core due to intense thermal stimulation of the body and head and resultant peripheral vasoconstriction. Dorsal head and upper chest immersion in cold water increases the rate of core cooling and decreases potential survival time.

Emergency treatment of hypothermia.

This review considers several recent concepts regarding aetiology and treatment of accidental hypothermia. The importance and effectiveness of shivering heat production in the attenuation and reversal of hypothermia is described. Immediately following removal from cold stress, the patient is in danger of a deteriorating condition that may be due to collapse of arterial pressure and/or continued decrease of core temperature. Several controversies are discussed. It is advised that, when possible, patients should be actively but gently warmed as soon as possible (especially if arrival at the emergency department will take greater than 45 min). Extra time should be taken to check for life signs before cardiopulmonary resuscitation is initiated. Chest compressions should proceed at regular normothermic rates and care should be taken to not overventilate the patient. In the emergency department, several factors should be considered before deciding on a treatment regimen. These factors include level of consciousness, cardiovascular stability, core temperature and the direction of change of core temperature. It may be advantageous to transport the more severely hypothermic patient to a more advanced care facility even though transport time may be greater.

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).

Contribution of shivering in leg muscles to heat production in Japanese quail

We estimated heat production in Japanese quail (Coturnix coturnix japonica) by measuring oxygen uptake using open-circuit respirometry as ambient temperature was decreased gradually from 26 to 3.5 °C. At the same time, the intensity of shivering was estimated in both the leg muscles and the flight muscles by measuring electromyograms. Metabolic heat production increased in a linear fashion as ambient temperature decreased. Shivering intensity increased at the same linear rate in the leg muscles as in the flight muscles as ambient temperature decreased. The leg muscles produce a substantial fraction (about 1/4) of the total shivering heat production at low ambient temperatures. Shivering occurred in bursts; the onset of a burst in the leg muscles was precisely synchronized with the onset of a burst in the flight muscles.

Sources of heat production in animals emerging from hibernation

Experiments on susliks showed that the active warming of these animals is closely linked with intensification of shivering heat production at all stages of emergence from hibernation. Shivering heat production by the proximal muscles plays the predominant role in this case.

Spinal cord thermosensitivity and sorting of neural signals in cold-exposed rats.

In the present study, data relevant to the presence or absence of sorting of neural signals were obtained by evaluating the thermal responses to spinal warming in the chronically prepared rat. Specifically, shivering activity and the rate of oxygen consumption (VO2) were measured in unanesthetized rats during cold exposure (10-16 degrees C). Warming the spinal cord at the level of T2 resulted in a significant decrease in shivering (P less than 0.001), without a significant change in VO2. The shivering response was reversed upon cessation of heating. These results are interpreted as indicating a direct influence of spinal cord temperature on shivering but not nonshivering thermogenesis in the rat. Similarly, in previous work with the rat, we have obtained data supporting hypothalamic receptor control of nonshivering but not shivering heat production. These findings are thus consistent with the suggestion that in the rat there occurs a sorting of neural signals. That is, impulses from the three thermoreceptor locations are not integrated in an identical manner for the control of shivering and nonshivering thermogenesis.

Shivering and nonshivering thermogenesis in the bat (Myotis myotis Borkh.) during arousal from hibernation

Nonshivering thermogenesis exists in the bat (Myotis myotis Borkh.) arousing from hibernation at environmental temperatures of 4–6 °C. Nonshivering thermogenesis is essential for the start of the arousal, and it is stimulated by noradrenaline since hexamethonium prevents the increase in metabolism and body temperature. Injection of noradrenaline abolishes this inhibition by hexamethonium by inducing nonshivering thermogenesis. After simultaneous administration of hexamethonium and alderlin no calorigenic effect of noradrenaline occurs. Shivering heat production during arousal appears at body temperatures between 10 and 17 °C predominantly. In normothermic bats the calorigenic effect of noradrenaline was observed, which indicates that nonshivering thermogenesis might also be present in awake animals. During arousal at 25 °C a great increase in intensity of shivering was observed. Elimination of nonshivering thermogenesis by hexamethonium does not prevent the attainment of the homoiothermic level of body temperature, and administration of noradrenaline does not speed up the process of arousal. As is evident from the metabolic capacity of the brown fat, the heat derived from this organ could maximally participate in total metabolism by 25% at the beginning and at the late period of arousal. In the middle range of body temperatures its significance for total metabolism is only about 10–13%.