• K. Cena
  • J. A. Clark
  • J. R. Spotila


Environmental physiologists have devoted many years of study to the mechanisms of thermoregulation, but the majority of research effort in this field has been concentrated on the physiology of the control of body temperature, with emphasis on various biochemical processes involved. However, the mandatory requirement for effective thermoregulation is not only possession of a thermostat, but also the existence of an interfacing structure that can modify the flow of heat between the body core and the environment. In homeothermic birds and mammals most body heat is produced internally (endothermy) and possession of adequate insulation to retain that heat is usually provided by the skin and hair coat or feathers. In poikilothermic amphibians and reptiles most body heat is absorbed from the environment (ectothermy) and the skin serves to control rates of heat and water exchange such that favorable body temperatures are maintained. Despite the fact that the physics of heat transfer through animal insulators is not mathematically complex, it has, until recently (Mitchell 1977, Cena and Clark 1979), received less attention than a number of secondary subcutaneous mechanisms concerned in thermoregulation and in homeostasis in particular. The aim of this chapter is to review current knowledge of the biophysics of animal insulators, and to describe how their structure controls the processes of energy exchange between the internal and ambient environments.


Heat Transfer Heat Loss Evaporative Cool Evaporative Water Loss Evaporative Heat Loss 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Allen TE, Bennett JW, Donegan SM, Hutchinson JCD (1970) Moisture, its accumulation and site of evaporation in the coats of sweating cattle. J Agric Sci 74: 247–258CrossRefGoogle Scholar
  2. Bakken GS, Gates DM (1974) Linearized heat transfer relations in biology. Science 183: 976–977PubMedCrossRefGoogle Scholar
  3. Bartholomew GA (1982) Physiological control of body temperature. In: Gans C, Pough FH (eds) Biology of the reptilia, vol XII. Academic Press, London New York, pp 167–211Google Scholar
  4. Bartholomew GA, Lasiewski RC (1965) Heating and cooling rates, heart rate and simulated diving in the Galapagos marine iguana. Comp Biochem Physiol 16: 573–582PubMedCrossRefGoogle Scholar
  5. Bennett JW (1964) Thermal insulation of cattle coats. Proc Aust Soc Anim Prod 5: 160–166Google Scholar
  6. Calder WA, King JR (1963) Evaporative cooling in the Zebra Finch. Experienta 19: 603, 1–3Google Scholar
  7. Campbell GS (1977) An introduction to environmental biophysics. Springer, Berlin Heidelberg New YorkCrossRefGoogle Scholar
  8. Campbell GS, McArthur AJ, Monteith JL (1980) Wind-speed dependence of heat and mass transfer through coats and clothing. Boundary-Layer Meteorol 18: 485–493CrossRefGoogle Scholar
  9. Cena K (1974) Radiative heat loss from animals and man. In: Monteith JL, Mount LE (eds) Heat loss from animals and man. Butterworths, London, pp 33–58Google Scholar
  10. Cena K, Clark JA (1973) Thermal radiation from animal coats: coat structure and measurements of radiative temperature. Phys Med Biol 18: 432–443PubMedCrossRefGoogle Scholar
  11. Cena K, Clark JA (1978) Thermal resistance units. J Therm Biol 3: 173–174CrossRefGoogle Scholar
  12. Cena K, Clark JA (1979) Transfer of heat through animal coats and clothing. In: Robertshaw D (ed) International review of physiology. Environmental physiology Ill, vol XX. Univ Park Press, Baltimore, pp 1–42Google Scholar
  13. Cena K, Clark JA (1981) Bioengineering, thermal physiology and comfort. Elsevier, Amsterdam Oxford New YorkGoogle Scholar
  14. Cena K, Monteith JL (1975a) Transfer processes in animal coats. I. Radiative transfer. Proc R Soc Lond [Biol] 188: 377–393Google Scholar
  15. Cena K, Monteith JL (1975b) Transfer processes in animal coats. II. Conduction and convection. Proc R Soc Lond [Biol] 188: 395–411Google Scholar
  16. Cena K, Monteith JL (1975e) Transfer processes in animal coats. III. Water vapour diffusion. Proc R Soc Lond [Biol] 188: 413–423Google Scholar
  17. Clark JA, MacLeod GD, Charles DR (1982) Causes of feather wear in poultry. Livestock environment I I. ASAE, St Joseph, MichGoogle Scholar
  18. Dawson TJ, Brown GD (1970) A comparison of the insulative and reflective properties of the fur of desert kangaroos. Comp Biochem Physiol 37: 23–38CrossRefGoogle Scholar
  19. Dawson WR, Bartholomew GA, Bennett AF (1977) A reappraisal of the aquatic specializations of the Galapagos marine iguana (Amblyrhynchus cristatus). Evolution 31: 891–897CrossRefGoogle Scholar
  20. Drane CR (1981) The thermal conductivity of the skin of crocodilians. Comp Biochem Physiol 68A: 107–110CrossRefGoogle Scholar
  21. Drane CR, Webb GJW (1980) Functional morphology of the dermal vascular system of the Australian lizard Tiliqua scincoides. Herpetologica 36: 60–66Google Scholar
  22. Evand KE, Moen AN (1975) Thermal exchange between Sharp-Tailed Grouse and their winter environment. Condor 77: 160–168CrossRefGoogle Scholar
  23. Eyal E (1963) Shorn and unshorn Awassi sheep. J Agric Sci 60: 159–193CrossRefGoogle Scholar
  24. Gatenby RM, Monteith JL, Clark JA ( 1983 a) Temperature and humidity gradients in a sheep’s fleece I. Gradients in steady state. Agric Meteorol 29: 1–10Google Scholar
  25. Gatenby RM, Monteith JL, Clark JA ( 1983 b) Temperature and humidity gradients in a sheep’s fleece II. The energetic significance of transients. Agric Meteorol 29: 83–101Google Scholar
  26. Gates DM (1973) Energy exchange in the biosphere. Harper and Row, New YorkGoogle Scholar
  27. Gates DM, Schmerl RB (1975) Perspectives of biophysical ecology. Springer, Berlin Heidelberg New YorkCrossRefGoogle Scholar
  28. Gebremedhind KG, Porter WP (1983) Sensitivity analysis of heat flow through irradiated fur of calves. Proc Am Soc Agric Eng Tech Pap NAR: 83–406Google Scholar
  29. Grigg GC, Alchin J (1976) The role of the cardiovascular system in thermo-regulation of Crocodylusjohnstoni. Physiol Zool 49: 24–36Google Scholar
  30. Hammel HT (1955) Thermal properties of fur. Am J Physiol 182: 369–376PubMedGoogle Scholar
  31. Heinrich B (1976) Heat exchange in relation to blood flow between thorax and abdomen in bumblebees. J Exp Biol 64: 561–566PubMedGoogle Scholar
  32. Johnson KG (1971) The discharge of sweat in Welsh mountain sheep. J Physiol (Lond) 215: 743Google Scholar
  33. King JR (1963) Oxygen consumption and body temperature in relation to ambient temperature in the White-Crowned Sparrow. Comp Biochem Physiol 12: 13–24Google Scholar
  34. King JR (1973) Energetics of reproduction in birds. In: Breeding biology of birds. Nall Acad Sci, WashingtonGoogle Scholar
  35. Kooyman GL, Gentry RL, Bergman WP, Hammel HT (1976) Heat loss in Penguins during immersion and compression. Comp Biochem Physiol [A] 54: 75–80CrossRefGoogle Scholar
  36. Kowalski GJ, Mitchell JW (1980) An experimental investigation of the convective heat transfer mechanisms within a fur layer. Am Soc Mech Eng Tech Pap 80-WA/HT28Google Scholar
  37. Lentz CP, Hart JS (1960) The effect of wind and moisture on heat loss through the fur of newborn caribou. Can J Zool 38: 679–688CrossRefGoogle Scholar
  38. Lillywhite HB, Maderson PFA (1982) Skin structure and permeability. In: Gans C, Pough FH (eds) Biology of the Reptilia, vol XII. Academic Press, London New YorkGoogle Scholar
  39. Lucas AM, Stettenheim PR (1982) Avian anatomy, integument, USDA Agric Handb 362, Washington DCGoogle Scholar
  40. Lundy H (1978) Recent studies in poultry calorimetry. Proc Nutr Soc 37: 55–63PubMedCrossRefGoogle Scholar
  41. Lustick S (1969) Bird energetics: effects of artificial radiation. Science 163: 387–390PubMedCrossRefGoogle Scholar
  42. MacLeod GD (1980) Mechanical properties of contour feathers. J Exp Biol 87: 65–71Google Scholar
  43. MacLeod GD, Clark JA (1980) An index of feather cover. J Agric Sci 95: 21–24CrossRefGoogle Scholar
  44. Mahoney SM, King JR (1977) The use of equivalent black-body temperature in the thermal energetics of small birds. J Therm Biol 2: 115–120CrossRefGoogle Scholar
  45. McArthur AJ, Monteith JL (1980) Air movement and heat loss from sheep. II. Thermal insulation of fleece in wind. Proc R Soc Lond [Biol] 209: 209–217Google Scholar
  46. McNeil-Alexander R (1971) Size and shape. Arnold, LondonGoogle Scholar
  47. Misson BH (1977) The relationships between body temperature and metabolic rate in the neonate fowl. J Therm Biol 2: 107–110CrossRefGoogle Scholar
  48. Mitchell D (1974) Physical basis of thermoregulation. In: Robertshaw D (ed) International review of physiology. Environmental physiology I, vol VII. Butterworths, London, pp 1–32Google Scholar
  49. Mitchell D (1977) Physical basis of thermoregulation. In: Robertshaw D (ed) International review of physiology. Environmental physiology II, vol XV. Univ Park Press, Baltimore, pp 1–27Google Scholar
  50. Mitchell MA (1985a) The effects of air velocity on convective and radiant heat transfer from domestic fowls. Br Poult Sci 26: 413–423PubMedCrossRefGoogle Scholar
  51. Mitchell MA (1985 b) Measurement of convective heat transfer in birds; a wind tunnel calorimeter. J Therm Biol 10:87–95Google Scholar
  52. Monteith JL (1973) Principles of environmental physics. Arnold, LondonGoogle Scholar
  53. Monteith JL, Mount LE (1974) Heat loss from animals and man. Butterworths, LondonGoogle Scholar
  54. Moote I (1955) The thermal insulation of caribou pelts. Text Res J 25: 832–834CrossRefGoogle Scholar
  55. Morgareidge KR, White FN (1969) Cutaneous vascular changes during heating and cooling in the Galapagos marine iguana. Nature 223: 587–591PubMedCrossRefGoogle Scholar
  56. Mount LE (1974) The concept of thermal neutrality. In: Monteith JL, Mount LE (eds) Heat loss from animals and man. Butterworths, London, pp 425–439Google Scholar
  57. Mount LE (1979) Adaptation to thermal environment. Butterworths, LondonGoogle Scholar
  58. Norris KS (1967) Color adaptation in desert reptiles and its thermal relationships. In: Milstead WW (ed) Lizard ecology: a symposium. Univ Missouri Press, Columbia, Mo, pp 162–229Google Scholar
  59. O’Neill SJB, Balnave D, Jackson N (1971) The influence of feathering and environmental temperature on the heat production and efficiency of utilization of metabolizable energy by the mature cockerel. J Agric Sci 77: 293–305CrossRefGoogle Scholar
  60. Pinshaw B, Fedak MA, Battles DR, Schmidt-Nielsen K (1976) Energy expenditure for locomotion and thermoregulation in Emperor Penguins. Am J Physiol 231: 903–912Google Scholar
  61. Poczopko P (1972) Thermal insulation in goslings. Acta Physiol Pol 23: 834–851Google Scholar
  62. Porter WP (1967) Solar radiation through the living body walls of vertebrates with emphasis on desert reptiles. Ecol Monogr 37: 273–296CrossRefGoogle Scholar
  63. Porter WP, Gates DM (1969) Thermodynamic equilibria of animals with environment. Ecol Monogr 39: 245–270CrossRefGoogle Scholar
  64. Porter WP, Norris KS (1969) Lizard reflectivity change and its effect on light transmiss’.on through body wall. Science 163: 482–484PubMedCrossRefGoogle Scholar
  65. Richards SA (1971) The significance of changes in the temperature of the skin and body core of the chicken in the regulation of heat loss. J Physiol 216: 1–10PubMedGoogle Scholar
  66. Richards SA (1976) Evaporative water loss in domestic fowls and its partition in relation to ambient temperature. J Agric Sci 87: 527–532CrossRefGoogle Scholar
  67. Robinson DE, Campbell GS, King JR (1976) The evaluation of heat exchange in small birds. J Comp Physiol 105: 153–166Google Scholar
  68. Roller WL, Dale AC (1963) Heat losses from Leghorn layers at warm temperatures. Trans ASAE 6: 136–139Google Scholar
  69. Scholander PF, Walters V, Hock R, Irving L (1950) Body insulation of some Arctic and tropical mammals and birds. Biol Bull 99: 225–236PubMedCrossRefGoogle Scholar
  70. Smith EN, Robertson SL, Davies DG (1978) Cutaneous blood flow during heating and cooling in the American alligator. Am J Physiol 235: R160 - R167PubMedGoogle Scholar
  71. Spitila JR (1980) Constraints of body size and environment on the temperature regulation of dinosaurs. In: Thomas RDK, Olson EC (eds) A cold look at the warm-blooded dinosaurs. AAAS Symp, vol XXVIII. Westview Press, Boulder, p 233–254Google Scholar
  72. Spotila JR, Berman E (1976) The role of the skin in the evaporative water loss from amphibians and reptiles. Comp Biochem Physiol [A] 55: 407–411CrossRefGoogle Scholar
  73. Spotila JR, Gates DM (1975) Body size, insulation and optimum body temperatures of homeotherms. In: Gates DM, Schmerl RB (eds) Perspectives of biophysical ecology. Springer, Berlin Heidelberg New York, pp 291–301CrossRefGoogle Scholar
  74. Spotila JR, Soule OH, Gates DM (1972) The biophysical ecology of the alligator, energy budgets and climate spaces. Ecology 43: 1094–1102CrossRefGoogle Scholar
  75. Taylor GR, Dmi’el R, Fedak M, Schmidt-Nielsen K (1971) The energetic cost of running and heat balance in a large bird — the Rhea. Am J Physiol 221: 597–601PubMedGoogle Scholar
  76. Tracy CR (1972) Newton’s Law: its application for expressing heat losses from homeotherms. Bioscience 22: 656–659CrossRefGoogle Scholar
  77. Tracy CR (1976) A model of the dynamic exchanges of water and energy between a terrestrial amphibian and its environment. Ecol Monogr 46: 293–326CrossRefGoogle Scholar
  78. Tracy CR (1982) Biophysical modelling in reptilian physiology and ecology. In: Gans C, Pough FH (eds) Biology of the reptilia, vol XII. Academic Press, London New York, pp 275–320Google Scholar
  79. Tregear RT (1965) Hair density, windspeed and heat loss in mammals. J Appl Physiol 20: 796–801PubMedGoogle Scholar
  80. VanDilla M, Day R, Siple PA (1968) Special problems of hands. In: Newburgh LH (ed) Physiology of heat regulation and the science of clothing. Hafner, New YorkGoogle Scholar
  81. Van Kampen M (1976) Activity and energy expenditure in laying hens, I, II and III. J Agric Sci 86:471–473 and 87:81–84, 85–88Google Scholar
  82. Walsberg GE (1980) The glossy appearance of a black bird: Thermal effects. J Therm Biol 5: 185–187CrossRefGoogle Scholar
  83. Walsberg GE (1982) Coat color, solar heat gain, and conspicuousness in the Phainopepla. Auk 99: 495–502Google Scholar
  84. Walsberg GE, King JR (1978a) The energetic consequences of incubation for two passerine species. Auk 95: 644–655Google Scholar
  85. Walsberg GE, King JR (1978b) The relationship of the external surface area of birds to skin surface area and body mass. J Exp Biol 76: 185–189Google Scholar
  86. Walsberg GE, Campbell GS, King JR (1978) Animal coat color and radiative heat gain: a re-evaluation. J Comp Physiol 126: 211–222Google Scholar
  87. Walton HK, Dale AC (1963) Radiant, convective and latent heat losses from mature White Leghorn chickens. Trans ASAE 6: 15–25Google Scholar
  88. Wathes CM, Clark JA (1981 a) Sensible heat transfer from the fowl: boundary layer resistance of a model fowl. Br Poult Sci 22: 161–173Google Scholar
  89. Wathes CM, Clark JA (1981 b) Sensible heat transfer from the fowl: thermal resistance of the pelt. Br Poult Sci 22: 175–183Google Scholar
  90. Wathes CM, Clark JA (1981c) Sensible heat transfer from the fowl: radiative and convective heat losses from a flock of broiler chickens. Br Poult Sci 22: 185–196PubMedCrossRefGoogle Scholar
  91. Webster MD, Campbell GS, King JR (1984) Cutaneous resistance to water vapor diffusion in pigeons and the role of the plumage. Physiol Zool (in press)Google Scholar
  92. Weinheimer CJ, Pendergast DR, Spotila JR, Wilson DR, Standora EA (1982) Peripheral circulation in Alligator mississippiensis: Effects of diving, fear, movement, investigator activities, and temperature. J Comp Physiol 148: 57–63Google Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 1986

Authors and Affiliations

  • K. Cena
    • 1
  • J. A. Clark
    • 2
  • J. R. Spotila
    • 3
  1. 1.Occupational Health ProgramMcMaster University Medical CenterHamiltonCanada
  2. 2.Department of Physiology and Environmental StudiesUniversity of Nottingham School of AgricultureSutton Bonington, Loughborough, Leics.England
  3. 3.Department of BiologyState University College at BuffaloBuffaloUSA

Personalised recommendations