Human physiological responses to cold exposure.

Thermal energy is transferred within and between bodies via several avenues, but for most unprotected human cold exposures, particularly during immersion, convective heat loss dominates. Lower tissue temperatures stimulate thermoreceptors, and the resultant afferent flow elicits autonomic homoeostatic responses (thermogenesis and vasoconstriction) that regulate body temperature within a narrow range. The most powerful effector responses occur when both superficial and deep thermoreceptors are cooled simultaneously, but thermoeffector activation can also occur as a result of peripheral cooling alone. The responses to cold, and the hazards associated with cold exposure, are moderated by factors which influence heat production and heat loss, including the severity and duration of cold stimuli, accompanying exercise, the magnitude of the metabolic response, and individual characteristics such as body composition, age, and gender. Cold stress can quickly overwhelm human thermoregulation with consequences ranging from impaired performance to death. This review provides a comprehensive overview of the human physiological responses to acute cold exposure.

[1]  Prescott Lf,et al.  Accidental hypothermia. A common condition. , 1962 .

[2]  S. Robinson,et al.  TEMPERATURE REGULATION IN EXERCISE. , 1963, Pediatrics.

[3]  G. W. Molnar,et al.  Exchanges of heat and tolerances to cold in men exposed to outdoor weather. , 1946, The American journal of physiology.

[4]  A. Jenkins,et al.  Interactions between temperature and human leptin physiology in vivo and in vitro , 2004, European Journal of Applied Physiology.

[5]  I B Mekjavic,et al.  The role of shivering thermogenesis and total body insulation in core cooling rate. , 1987, The Annals of physiological anthropology = Seiri Jinruigaku Kenkyukai kaishi.

[6]  O. Edholm,et al.  THE PHYSIOLOGY OF CHANNEL SWIMMERS , 1955 .

[7]  T. J. Doubt,et al.  Prolonged whole-body cold water immersion: fluid and ion shifts. , 1989, Journal of applied physiology.

[8]  M. Tipton,et al.  Supraventricular arrhythmias following breath-hold submersions in cold water. , 1994, Undersea & hyperbaric medicine : journal of the Undersea and Hyperbaric Medical Society, Inc.

[9]  R. E. Sloan,et al.  Cooling rates of young people swimming in cold water. , 1973, Journal of applied physiology.

[10]  M. Sawka,et al.  Human vascular fluid responses to cold stress are not altered by cold acclimation. , 1987, Undersea biomedical research.

[11]  S. Horvath,et al.  Compensatory cardiovascular responses during an environmental cold stress, 5 degrees C. , 1970, Journal of applied physiology.

[12]  L. Pugh The influence of wind resistance in running and walking and the mechanical efficiency of work against horizontal or vertical forces , 1971, The Journal of physiology.

[13]  I Holmér,et al.  Metabolic and thermal response to swimming in water at varying temperatures. , 1974, Journal of applied physiology.

[14]  C. Wilson Climatology of the cold regions : Northern Hemisphere II , 1969 .

[15]  A. C. Burton THE RANGE AND VARIABILITY OF THE BLOOD FLOW IN THE HUMAN FINGERS AND THE VASOMOTOR REGULATION OF BODY TEMPERATURE , 1939 .

[16]  W. Krone,et al.  Rapid rise in plasma glucagon induced by acute cold exposure in man and rat , 2004, Pflügers Archiv.

[17]  K B Pandolf,et al.  Thermal responses during arm and leg and combined arm-leg exercise in water. , 1984, Journal of applied physiology: respiratory, environmental and exercise physiology.

[18]  M. Eriksen,et al.  Effect of local warming on hand and finger artery blood velocities. , 1995, The American journal of physiology.

[19]  B. Ekblom,et al.  Influence of muscle temperature on maximal muscle strength and power output in human skeletal muscles. , 1979, Acta physiologica Scandinavica.

[20]  I. Macdonald,et al.  Physiological responses to a cold, wet, and windy environment during prolonged intermittent walking. , 1997, The American journal of physiology.

[21]  J. Hayward,et al.  Wet-cold exposure and hypothermia: thermal and metabolic responses to prolonged exercise in rain. , 1996, Journal of applied physiology.

[22]  I. Mekjavic,et al.  Effect of hypoglycemia on thermoregulatory responses. , 1996, Journal of applied physiology.

[23]  N. Taylor,et al.  Preferred room temperature of young vs aged males: the influence of thermal sensation, thermal comfort, and affect. , 1995, The journals of gerontology. Series A, Biological sciences and medical sciences.

[24]  I. Korhonen,et al.  Serum levels of thyroid and adrenal hormones, testosterone, TSH, LH, GH and prolactin in men after a 2-h stay in a cold room. , 1988, Acta physiologica Scandinavica.

[25]  S. Lennquist Cold-induced diuresis. A study with special reference to electrolyte excretion, osmolal balance and hormonal changes. , 1972, Scandinavian journal of urology and nephrology.

[26]  M. Vincent,et al.  The effects of cold immersion and hand protection on grip strength. , 1988, Aviation, space, and environmental medicine.

[27]  P. Tikuisis,et al.  Physiological responses of exercised-fatigued individuals exposed to wet-cold conditions. , 1999, Journal of applied physiology.

[28]  J. Cotter,et al.  Epidemiology of hypothermia: fatalities and hospitalisations in New Zealand. , 1994, Australian and New Zealand journal of medicine.

[29]  G. W. Molnar,et al.  Survival of hypothermia by men immersed in the ocean. , 1946, Journal of the American Medical Association.

[30]  Golden Fs Proceedings: Recognition and treatment of immersion hypothermia. , 1973 .

[31]  D. Kenshalo Changes in the cool threshold associated with phases of the menstrual cycle. , 1966, Journal of applied physiology.

[32]  L. Rowell Human Circulation: Regulation During Physical Stress , 1986 .

[33]  B. Saltin,et al.  Maximal oxygen uptake and heart rate in various types of muscular activity. , 1961, Journal of applied physiology.

[34]  W. Keatinge,et al.  The effect of repeated daily exposure to cold and of improved physical fitness on the metabolic and vascular response to cold air , 1961, The Journal of physiology.

[35]  H. Hatfield,et al.  Thermal Conductivity of Human Fat and Muscle , 1951, Nature.

[36]  I. Mekjavic,et al.  Contribution of core cooling rate to shivering thermogenesis during cold water immersion. , 1991, Aviation, space, and environmental medicine.

[37]  Nunneley Sa,et al.  Physiological responses of women to thermal stress: a review. , 1978 .

[38]  M. Campbell,et al.  Sudden infant deaths and cold weather: was the rise in infant mortality in 1986 in England and Wales due to the weather? , 1991, Paediatric and perinatal epidemiology.

[39]  W. L. Kenney,et al.  Methylcholine-activated eccrine sweat gland density and output as a function of age. , 1988, Journal of applied physiology.

[40]  A. Jenkins,et al.  Cold-water acclimation does not modify whole-body fluid regulation during subsequent cold-water immersion , 2004, European Journal of Applied Physiology.

[41]  P. Lomax,et al.  Body temperature : regulation, drug effects, and therapeutic implications , 1979 .

[42]  L. A. Stephenson,et al.  Thermoregulation in Women , 1993, Exercise and sport sciences reviews.

[43]  A N Exton-Smith,et al.  Effects of age on body temperature and blood pressure in cold environments. , 1985, Clinical science.

[44]  Mikael Gennser,et al.  Immersion deaths and deterioration in swimming performance in cold water , 1999, The Lancet.

[45]  Hardy Jd Control of heat loss and heat production in physiologic temperature regulation. , 1953 .

[46]  M. Cabanac,et al.  Evidence form behavioral thermoregulatory responses of a shift in setpoint temperature related to the menstrual cycle. , 1971, Journal de physiologie.

[47]  W R Keatinge,et al.  Failure of thermoregulation in the cold during hypoglycaemia induced by exercise and ethanol , 1973, The Journal of physiology.

[48]  H. H. Mitchell,et al.  Shivering and heat production in men exposed to intense cold. , 1967, Journal of applied physiology.

[49]  J. Shepherd,et al.  Responses of the Superficial Limb Veins of the Dog to Changes in Temperature , 1968, Circulation research.

[50]  M. Allwood,et al.  Muscle and skin blood flow in the human forearm during insulin hypoglycaemia , 1959, The Journal of physiology.

[51]  P. Payne,et al.  Environmental Conditions and Body Temperatures of Elderly Women Living Alone or in Local Authority Home , 1971, British medical journal.

[52]  I. Jacobs,et al.  Thermoregulatory Thermogenesis in Humans during Cold Stress , 1994, Exercise and sport sciences reviews.

[53]  G. Schroth,et al.  Outcome of Survivors of Accidental Deep Hypothermia and Circulatory Arrest Treated with Extracorporeal Blood Warming , 1997, The New England journal of medicine.

[54]  M L Collis,et al.  Thermal balance and survival time prediction of man in cold water. , 1975, Canadian journal of physiology and pharmacology.

[55]  Horvath Sm,et al.  Thermoregulation in surfers and nonsurfers immersed in cold water. , 1978 .

[56]  O G Edholm,et al.  Temperature and blood flow in the human forearm , 1946, The Journal of physiology.

[57]  A N Exton-Smith,et al.  Urban hypothermia: preferred temperature and thermal perception in old age. , 1981, British medical journal.

[58]  S. Horvath,et al.  Adaptations in man's adrenal function in response to acute cold stress. , 1974, Journal of applied physiology.

[59]  I. Mekjavic,et al.  Human temperature regulation during narcosis induced by inhalation of 30% nitrous oxide , 1992 .

[60]  James M. Tanner,et al.  Fetus into Man , 1978 .

[61]  H. Pekkarinen,et al.  Different thermal sensitivity to exercise and cold in men and women , 1999 .

[62]  The cutaneous microcirculation of the forearm in young and old subjects. , 1991, Microvascular research.

[63]  E. Buskirk,et al.  Effects of physical conditioning in man on thermal responses to cold air , 1972, International journal of biometeorology.

[64]  R H Fox,et al.  A comparison of thermoregulatory function in men and women. , 1968, The Journal of physiology.

[65]  S. Horvath,et al.  Urinary responses to cold temperature during water immersion. , 1985, The American journal of physiology.

[66]  I. Roddie Sympathetic vasodilatation in human skin. , 2003, The Journal of physiology.

[67]  M. Sawka,et al.  Thermoregulation during cold exposure: effects of prior exercise. , 1999, Journal of applied physiology.

[68]  J. Frim,et al.  Effects of endurance fitness on responses to cold water immersion. , 1984, Aviation, space, and environmental medicine.

[69]  Hong Sk,et al.  Comparison of diving and nondiving women of Korea. , 1963 .

[70]  P. Auerbach Reviews and Notes: Emergency Medicine: Wilderness Medicine: Management of Wilderness and Environmental Emergencies , 1995, Annals of Internal Medicine.

[71]  J D Hardy,et al.  Differences between Men and Women in Their Response to Heat and Cold. , 1940, Proceedings of the National Academy of Sciences of the United States of America.

[72]  C. Basbaum Induced hypothermia in peripheral nerve: Electron microscopic and electrophysiological observations , 1973, Journal of neurocytology.

[73]  J. House,et al.  Work Enhancement and Thermal Changes during Intermittent Work in Cool Water after Carbohydrate Loading , 1990 .

[74]  C. Wade,et al.  Regional heat loss in resting man during immersion in 25.2 degrees C water. , 1979, Aviation, space, and environmental medicine.

[75]  L. G. C. PUGH,et al.  Clothing Insulation and Accidental Hypothermia in Youth , 1966, Nature.

[76]  D. E. Bass,et al.  Responses of body fluid compartments to heat and cold. , 1956, Physiological reviews.

[77]  B. Brodie,et al.  The sympathetic nervous system as a homeostatic mechanism. I. Absolute need for sympathetic nervous function in body temperature maintenance of cold-exposed rats. , 1967, The Journal of pharmacology and experimental therapeutics.

[78]  E. Nadel,et al.  Energy exchanges of swimming man. , 1974, Journal of applied physiology.

[79]  M. Epstein Renal effects of head-out water immersion in man: implications for an understanding of volume homeostasis. , 1978, Physiological reviews.

[80]  J A Stolwijk,et al.  Comparative thermoregulatory responses of resting men and women. , 1978, Journal of applied physiology: respiratory, environmental and exercise physiology.

[81]  S. K. Hong,et al.  Energetics of wet-suit diving in Korean women breath-hold divers. , 1983, Journal of applied physiology: respiratory, environmental and exercise physiology.

[82]  K. Hagbarth,et al.  Thermoregulatory and rhythm‐generating mechanisms governing the sudomotor and vasoconstrictor outflow in human cutaneous nerves. , 1980, The Journal of physiology.

[83]  I Holmér,et al.  Oxygen uptake during swimming in man. , 1972, Journal of applied physiology.

[84]  R. Bates,et al.  Defining the cold regions of the Northern Hemisphere , 1966 .

[85]  P. Baker,et al.  Relationship between skinfold thickness and body cooling for two hours at 15°C. , 1956 .

[86]  H. Hensel,et al.  The central control of shivering and non‐shivering thermogenesis in the rat. , 1978, The Journal of physiology.

[87]  P. Greenhaff,et al.  Physiological responses to moderate cold stress in man and the influence of prior prolonged exhaustive exercise , 1998, Experimental physiology.

[88]  J. Johnson,et al.  Nonthermoregulatory control of human skin blood flow. , 1986, Journal of applied physiology.

[89]  R. White,et al.  Terrestrial Mammals in Cold , 1989 .

[90]  A. R. Lind,et al.  The duration of sustained contractions of the human forearm at different muscle temperatures , 1958, The Journal of physiology.

[91]  C. Pagni,et al.  ELECTROENCEPHALOGRAPHIC MODIFICATIONS INDUCED BY MODERATE AND DEEP HYPOTHERMIA IN MAN. , 1964, Acta neurochirurgica. Supplement.

[92]  Male-female responses in various body temperatures during and following exercise in cold air. , 1986 .

[93]  L G Pugh,et al.  Rectal temperatures, weight losses, and sweat rates in marathon running. , 1967, Journal of applied physiology.

[94]  G. Hervey Proceedings: Physiological changes encountered in hypothermia. , 1973, Proceedings of the Royal Society of Medicine.

[95]  E. Hultman,et al.  A study of the glycogen metabolism during exercise in man. , 1967, Scandinavian journal of clinical and laboratory investigation.

[96]  R H Fox,et al.  Body Temperatures in the Elderly: A National Study of Physiological, Social, and Environmental Conditions , 1973, British medical journal.

[97]  O. Grosser Ueber arterio-venöse Anastomosen an den Extremitätenenden beim Menschen und den krallentragenden Säugethieren , 1902 .

[98]  B. Lilja,et al.  Hemodynamic changes in man during immersion with the head above water. , 1972, Aerospace medicine.

[99]  H Ue,et al.  Autonomic responsiveness to acute cold exposure in obese and non-obese young women , 1999, International Journal of Obesity.

[100]  P. A. Nilsen,et al.  Resuscitation from accidental hypothermia of 13·7°C with circulatory arrest , 2000, The Lancet.

[101]  J. Morrison,et al.  Accidental hypothermia: the effect of initial body temperatures and physique on the rate of rewarming. , 1980, Aviation, space, and environmental medicine.

[102]  E. Buskirk,et al.  Metabolic and thermal responses of women during cooling in water , 1974 .

[103]  J. Boulant,et al.  Effects of testosterone, estradiol, and temperature on neurons in preoptic tissue slices. , 1986, The American journal of physiology.

[104]  W. Kraemer,et al.  Menstrual status and plasma vasopressin, renin activity, and aldosterone exercise responses. , 1989, Journal of applied physiology.

[105]  M. Tipton,et al.  Substrate utilisation during exercise and shivering , 1997, European Journal of Applied Physiology and Occupational Physiology.

[106]  T. Nagasaka,et al.  Effects of human menstrual cycle on thermoregulatory vasodilation during exercise , 2004, European Journal of Applied Physiology and Occupational Physiology.

[107]  G. Ferretti,et al.  Superficial shell insulation in resting and exercising men in cold water. , 1982, Journal of applied physiology: respiratory, environmental and exercise physiology.

[108]  D. Rennie,et al.  Tissue heat transfer in water: lessons from the Korean divers. , 1988, Medicine and science in sports and exercise.

[109]  O. Edholm,et al.  Man, hot and cold , 1978 .

[110]  P. Daniel,et al.  Arteriovenous anastomoses in the human external ear. , 1956, Journal of anatomy.

[111]  J. Bittel,et al.  Physical fitness and thermoregulatory reactions in a cold environment in men. , 1988, Journal of applied physiology.

[112]  S. K. Hong,et al.  Physical insulation of Korean diving women. , 1962, Journal of applied physiology.

[113]  W. Keatinge Cold immersion and swimming. , 1972, Journal of the Royal Naval Medical Service.

[114]  V. Hessemer,et al.  Influence of menstrual cycle on shivering, skin blood flow, and sweating responses measured at night. , 1985, Journal of applied physiology.

[115]  W. D. McArdle,et al.  Thermal adjustment to cold-water exposure in resting men and women. , 1984, Journal of applied physiology: respiratory, environmental and exercise physiology.

[116]  R. F. Goldman,et al.  Heat production from shivering. , 1960, Journal of applied physiology.

[117]  J. P. Henry,et al.  Circulatory basis of fluid volume control. , 1963, Physiological reviews.

[118]  Hall Jf Prediction of tolerance in cold water and life raft exposures. , 1972 .

[119]  J. Bittel,et al.  Reversal of cold induced haemoconcentration , 2004, European Journal of Applied Physiology and Occupational Physiology.

[120]  O. Bar-or,et al.  Thermoregulation during rest and exercise in the cold in pre- and early pubescent boys and in young men. , 1992, Journal of applied physiology.

[121]  W. Keatinge,et al.  Increases in platelet and red cell counts, blood viscosity, and arterial pressure during mild surface cooling: factors in mortality from coronary and cerebral thrombosis in winter. , 1984, British medical journal.

[122]  Keatinge Wr Death after shipwreck. , 1965 .

[123]  W. Keatinge,et al.  Exceptional case of survival in cold water. , 1986, British medical journal.

[124]  W. Keatinge The effects of subcutaneous fat and of previous exposure to cold on the body temperature, peripheral blood flow and metabolic rate of men in cold water , 1960, The Journal of physiology.

[125]  S. Horvath,et al.  Cardiovascular reactions to cold exposures differ with age and gender. , 1985, Journal of applied physiology.

[126]  S. K. Hong,et al.  Cold tolerance of long-distance runners and swimmers in Hawaii , 1977, International journal of biometeorology.

[127]  A N Exton-Smith,et al.  Functional changes in autonomic nervous responses with ageing. , 1980, Age and ageing.

[128]  F. Depocas Biochemical changes in exposure and acclimation to cold environments. , 1961, British medical bulletin.

[129]  I. Rothchild,et al.  The effects of dosage, and of estrogen, androgen or salicylate administration on the degree of body temperature elevation induced by progesterone. , 1952, Endocrinology.

[130]  W. Blackard,et al.  Insulin secretion in hypothermic dogs. , 1967, The American journal of physiology.

[131]  P Tikuisis,et al.  In vivo thermal conductivity of the human forearm tissues. , 1991, Journal of applied physiology.

[132]  Park Ys,et al.  Decrease in body insulation with exercise in cool water. , 1984 .

[133]  M. Hayward,et al.  Roles of subcutaneous fat and thermoregulatory reflexes in determining ability to stabilize body temperature in water. , 1981, The Journal of physiology.

[134]  O H Gauer,et al.  The regulation of extracellular fluid volume. , 1970, Annual review of physiology.

[135]  L. Fleisher,et al.  Adrenergic, respiratory, and cardiovascular effects of core cooling in humans. , 1997, The American journal of physiology.

[136]  E. Deacon,et al.  Some Statistical and Climatological Aspects of the Incidence of the Sudden Infant Death Syndrome , 1979, Australian paediatric journal.

[137]  M. Tipton,et al.  The effects of warming by active and passive means on the subsequent responses to cold water immersion , 2004, European Journal of Applied Physiology and Occupational Physiology.

[138]  D. Johnson,et al.  Plasma norepinephrine responses of man in cold water. , 1977, Journal of applied physiology: respiratory, environmental and exercise physiology.

[139]  G. Giesbrecht,et al.  Cold stress, near drowning and accidental hypothermia: a review. , 2000, Aviation, space, and environmental medicine.

[140]  S. Lennquist,et al.  Fluid balance and physical work capacity in humans exposed to cold. , 1974, Archives of environmental health.

[141]  S. Horvath,et al.  Metabolic responses of old people to a cold environment. , 1955, Journal of applied physiology.

[142]  H. Rintamäki,et al.  Plasma levels of endothelin-1 and atrial natriuretic peptide in men during a 2-hour stay in a cold room. , 1991, Acta physiologica Scandinavica.

[143]  I. Jacobs,et al.  Muscle glycogen utilization during shivering thermogenesis in humans. , 1988, Journal of applied physiology.

[144]  S. Horvath,et al.  The influence of cold upon the efficiency of man. , 1947, The Journal of aviation medicine.

[145]  L. P. Herrington,et al.  Physiological effects of heat and cold. , 1947, Annual review of physiology.

[146]  Kim Jk,et al.  Peripheral blood flow and heat flux of Korean women divers. , 1969 .

[147]  H. Hensel Thermoreception and temperature regulation. , 1981, Monographs of the Physiological Society.

[148]  I. Macdonald,et al.  Hypoglycaemia, hypothermia and shivering in man. , 1981, Clinical science.

[149]  P Tikuisis,et al.  Exertional fatigue, sleep loss, and negative energy balance increase susceptibility to hypothermia. , 1998, Journal of applied physiology.

[150]  I. Jacobs,et al.  Free fatty acid availability and temperature regulation in cold water. , 1989, Journal of applied physiology.

[151]  I. Jacobs,et al.  Rates of energy substrates utilization during human cold exposure , 2006, European Journal of Applied Physiology and Occupational Physiology.

[152]  M. Tipton,et al.  The initial responses to cold-water immersion in man. , 1989, Clinical science.

[153]  L. Rowell,et al.  Human cardiovascular adjustments to exercise and thermal stress. , 1974, Physiological reviews.

[154]  R. F. Goldman,et al.  Role of air temperature and wind in the time necessary for a finger to freeze. , 1970, Journal of applied physiology.

[155]  M. Eriksen,et al.  Fluctuations in blood flow to acral skin in humans: connection with heart rate and blood pressure variability. , 1993, The Journal of physiology.

[156]  M. Sawka,et al.  Thermoregulation during cold water immersion is unimpaired by low muscle glycogen levels. , 1989, Journal of applied physiology.

[157]  G. Marion,et al.  Core body temperature in the elderly and factors which influence its measurement. , 1991, Gerontology.

[158]  M. Sawka,et al.  Effects of body mass and morphology on thermal responses in water. , 1986, Journal of applied physiology.

[159]  P Tikuisis,et al.  Shivering onset, metabolic response, and convective heat transfer during cold air exposure. , 1991, Journal of applied physiology.

[160]  F. Girling METABOLIC RESPONSE TO NUDE HUMAN SUBJECTS TO ACUTE EXPOSURE AT 10 DEGREES C. , 1964, Canadian journal of physiology and pharmacology.