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Glossary of Terms For Thermal Physiology
Glossary of Terms For Thermal Physiology
Communications concerning the glossary may be sent the term acclimatization to describe the adaptive
to the Chairman of the IUPS Thermal Commission, or changes that occur within an organism in response to
to the Publisher. changes in the natural climate.
Refs: Eagan, C. J., Federation Proc. 22: 930, 1963;
Acknowledgement Greenleaf, J.E., & C.J. Greenleaf, NASA Tech. Mem.
This revision was initiated by James Mercer during his X-62, 008, 1970; Hart, J.S., Rev. Can. Biol. 16: 133,
term as Chairman of the I.U.P.S. Commission for Ther- 1957.
mal Physiology. Thermal Commission member Jurgen
Werner was responsible for producing the first working Acclimatization: Physiological or behavioral changes
draft and further updating of the glossary followed occurring within the lifetime of an organism that reduce
from numerous additional suggestions for new entries the strain caused by stressful changes in the natural cli-
and modifications by the current commission members mate (e.g., seasonal or geographical). → acclimation.
and other consulted experts. In the revised glossary, 11 Note: What is termed acclimatization may also be
terms have been removed, 49 of the original entries specified as a particular phenotypic adaptation; there-
have been modified and the definitions of 37 new terms fore, the usefulness of the term may be questioned.
have been added, increasing the overall number of en-
tries to 479. Acute-phase response (APR): The multifactorial ste-
reotyped response of an organism to infection, injury or
James Mercer, Chairman trauma. The APR comprises changes in plasma con-
I.U.P.S. Commission for Thermal Physiology centrations of trace metals, liver proteins (so-called
Department of Medical Physiology “acute phase proteins”), hormones, intermediary me-
Institute of Medical Biology tabolism, neutrophilia, and “sickness behavior”, which
Faculty of Medicine includes the development of fever, loss of food appe-
University of Tromsø tite, increased sleep, decreased motor activity, and de-
N-9037 Tromsø. creased alertness. The acute-phase response (APR) is
Norway. considered an early, non-specific host-defence re-
sponse, and is triggered by the release of cytokines
Glossary such as interleukin-1 (IL-1), IL-6, and tumor necrosis
factor α (TNFα).
Absolute humidity. → Humidity, absolute. Adaptation: Changes that reduce the physiological
strain produced by stressful components of the total
Absorptance, total radiant. → Radiant absorptance, environment. This change may occur within the life-
total. time of an organism (phenotypic) or be the result of ge-
netic selection in a species or subspecies (genotypic).
Accidental hypothermia. → Hypothermia, acciden- Acclimation as defined in this Glossary relates to phe-
tal. notypic adaptations to specified climatic components.
In thermal physiology, the use of the term adaptation
Acclimation: Physiological or behavioral changes oc- does not require specification of the climatic compo-
curring within an organism, which reduces the strain or nents of the total environment to which the organism
enhances endurance of strain caused by experimentally adapts, but the most obvious component is often de-
induced stressful changes in particular climatic factors. noted (e.g., adaptation to heat). There are no distinct
Note: The terms acclimation and acclimatization are terms that relate genotypic adaptations to the climate
etymologically indistinguishable. Both words have or particular components of climate.
been assigned several and different meanings (Green- Note: In comparison to adaptation as defined in neuro-
leaf & Greenleaf, 1970). The most useful of the as- physiology, the adaptive processes in thermal physiol-
signed meanings, adopted here, would seem to be those ogy usually occur with larger time constants.
of Hart (1957) and Eagan (1963) who use the term ac-
climation to describe the adaptive changes that occur Adaptation, genetic. → Adaptation, genotypic.
within an organism in response to experimentally in-
duced changes in particular climatic factors such as Adaptation, genotypic: A genetically fixed condition
ambient temperature in a controlled environment, and of a species or subspecies, or its evolution, which
favours survival in a particular total environment. → some conditions) are also endogenous antipyretics.
adaptation. All, however, have distinct mechanisms of action.
ipose tissue with selected distribution. It differs from the heat transfer.
white adipose tissue both structurally and functionally; Note: The most common method of indirect calorim-
its colour is brownish (yellow to reddish), depending on etry is to measure the uptake of oxygen and/or the elim-
its content of (mitochondrial) cytochromes and of fat ination of carbon dioxide, and to convert these values to
which is distributed in multipledroplets (multilocular) an equivalent quantity of heat. The calorific equivalent
as compared to white fat cells (monolocular) and also of a litre of oxygen is approximately 20 kJ of heat.
on its dense vascularization. Brown adipose tissue ap-
pears to be restricted to mammalian species, especially Calorimetry, partitional: The estimation of any single
of smaller body size, including the hibernators (→ Hi- term in the body heat balance equation from direct
bernation), but is also present in newborns of larger measurements of all other terms in the equation during
species including humans. Regulatory heat production the steady state. Ref: Winslow, C.E.A., L.P. Herrington,
has been identified as its main but possibly not its only & A.P. Gagge, Am. J. Physiol. 116: 641, 1936.
function. Its heat-generating capacity relates to the rate
of fatty acid oxidation which is not constrained by cel- Cenothermy: The condition of a temperature regula-
lular ATP catabolism. This property correlates with the tor when its core temperature is within +1 SD of the
presence of a specialized component in the inner mito- range associated with the normal postabsorptive resting
chondrial membrane named UCP1 (the uncoupling condition of the species in the thermoneutral zone.
protein or thermogenin, which establishes a proton (Gk. koinos-common, shared; therme-heat) Synonym:
shunt bypassing oxidative phosphorylation). The activ- euthermy, normothermy.
ity of brown adipose tissue is controlled by the sympa- Note: Ceno (common) seems a more appropriate prefix
thetic nervous system through release of adrenaline. than eu (well, good).
Note: Chronic exposure to cold (→ nonshivering ther-
mogenesis) and to excess food intake (→ diet-induced Change in heat storage. → Heat storage, change in.
thermogenesis) enhance the heat-generating capacity
of brown adipose tissue by hypertrophy (including in- Chemical temperature regulation. → Temperature
crease in UCP1 amount) and hyperplasia. The total regulation, chemical.
process is referred to as recruitment. In accordance with
the sympathetic control of brown adipose tissue, the Circadian: Relating to the approximate 24-h periodic-
increase in metabolic energy transformation of an ity of a free running biological rhythm, or to the exact
animal in a thermoneutral environment after systemic 24-h periodicity of an environmentally synchronized
injection of noradrenaline is considered as an estimate biological rhythm that persists with an approximate
of the thermogenetic capacity of brown adipose tissue. 24-h periodicity when not environmentally synchro-
nized. (L. circa-about, approximately; dies-day)
Brown fat. → Brown adipose tissue.
clo: A unit to express the relative thermal insulation
Calorimetry: The measurement of heat. In thermal values of various clothing assemblies.
physiology, the measurement of the heat transfer be- 1 clo = 0.18°C·m2 ·h · kcal–1 = 0.155°C ·m2 ·W–1
tween a tissue, an organ, or an organism and its envi-
ronment. (L. calor-heat; Gk. metria-act of measuring) Note: The clo is a unit developed to express thermal
→ calorimetry, direct and calorimetry, indirect. insulation (→ thermal resistance) in practical terms
and represents the insulation provided by the normal
Calorimetry, direct: The direct physical measurement indoor clothing of a sedentary worker in comfortable
of heat, usually the rate of transfer of heat (→ heat indoor surroundings (Ref: Gagge, A.P., A.C. Burton, &
transfer) between a tissue, an organ, or an organism H.C. Bazett, Science 94: 428, 1941). The term is used in
and its environment. heating and ventilation engineering in the determina-
tion of environmental conditions for human comfort.
Calorimetry, indirect: The measurement of the rate of
transfer of a material involved in the transformation of Cold-blooded: The thermal state of an animal in which
chemical energy into heat between a tissue, an organ, or core temperature remains close to ambient tempera-
an organism and its environment. The process requires ture when subjected to a low ambient temperature.
the calculation of the heat transfer from an empirically Synonym: bradymetabolic, poikilothermic. Ant-
established relation between the material transfer and onym: warm-blooded.
Note: The existence of only a small temperature gradi- Comfort. → Thermal comfort.
ent between the organism and its environment results
from the low rate of metabolic heat production Conductance, thermal. → Thermal conductance.
(bradymetabolism) of cold-blooded animals relative
to the high rate of metabolic heat production (ta- Conductive heat transfer (K): The net rate of heat
chymetabolism) of warm-blooded animals. Thus, the transfer in a solid material or a non-moving gas or
terms bradymetabolic and tachymetabolic are pre- fluid (i.e., by conduction) down a thermal gradient,
ferred to the terms cold-blooded and warm-blooded, within an organism, or between an organism and its ex-
because the first pair of terms relates to a more basic ternal environment. The latter is usually expressed in
physiological distinction and because the second pair of terms of unit area of the total body surface, i.e. as a heat
terms has been used with various meanings not all of flux, and is the quantity (K) in the body heat balance
which are entirely consistent with the definitions given equation in which (–K) = heat gain, and (+K) = heat
here. Since their core temperatures follow ambient loss. [W·m–2], [W], [W·m–3], [W·kg–1]
temperature, all cold-blooded animals are poikilother-
mic (i.e., many temperatured). Thus, cold-blooded, Conductive heat transfer coefficient. → Heat trans-
poikilothermic, and bradymetabolic are descriptions fer coefficient, conductive.
of related phenomena. The same cannot be said of
warm-blooded, homeothermic, and tachymetabolic Conductivity, thermal. →Thermal conductivity.
(→ warm-blooded).
Control: 1. the process by which a physiological vari-
Cold receptor. → Thermoreceptor. able becomes a function of information generated lo-
cally or transmitted from remote sources by neural or
Cold tolerance: The ability to tolerate low ambient hormonal signals (feedforward control); 2. the process
temperatures. This term comprises a variety of physi- by which a physiological system stabilizes (“regu-
ological properties. Certain homeotherms (tachymet- lates”) a variable, generally by means of an information
abolic species) are described as cold tolerant, because loop with negative feedback. The control loop com-
they are able to balance heat production and heat loss in prises the controlled (passive) system (e.g., the heat
the state of cenothermy at particularly low ambient transfer processes) and the controlling (active) system,
temperatures. This may be due to the development of which may be divided into sensor, processor and effec-
insulation or to the efficiency of metabolic heat pro- tor components.
duction and is often combined with an improved abil- Note: With the adoption of definition #2 from the ter-
ity to protect appendages from freezing by effective minology of technical control, further terms like
vascular control of local heterothermy. In homeo- set-point, controlled variable, load-error, nega-
therms, cold tolerance may also be achieved by gen- tive-feedback, servo-control, etc., have been introduced
eral heterothermy, including hibernation, i.e., by to describe the various components of the physiological
controlled forms of hypothermia. Poikilotherms processes of regulation. Control in the sense of defini-
(bradymetabolic species) are characterized as cold tion #2 and regulation are analogous terms in physiol-
tolerant, if they are able to survive low and even sub- ogy. However, the use of the term regulation should be
freezing body temperatures, either in the state of cryo- generally given preference in order to avoid confusion
thermy, or because intracellular formation of ice between the two definitions of control.
crystals is avoided during freezing. All forms of cold
tolerance may be subject to adaptation or accli- Control, proportional: A control concept with nega-
mat(izat)ion. tive feedback, obviously present in thermoregulation,
Note: While some types of what is termed cold toler- in which the controller activates thermoeffector mech-
ance have already been specified (cryothermy, hiber- anisms to an extent dependent on the deviation of the
nation), further specification and agreement upon controlled variable from its set-point. If the distur-
additional definitions seem necessary. bance (external or internal thermal load) persists, such a
control concept implies a load error, i.e., a permanent
Combined nonevaporative heat transfer coefficient. deviation of the controlled variable. However, such a
→ Heat transfer coefficient, combined nonevapora- deviation is much smaller than that which is present
tive. without feedback control.
Controlled temperature lability. → Temperature la- a section of the thermal core: however, in several
bility, controlled. mammalian and avian species its temperature may de-
viate to some extent from that of the remaining thermal
Convection, circulatory: Flow of blood in the vessels core due to selective brain cooling.
of the circulatory system. In thermal physiology, con-
vective transfer of heat with the blood stream from the Crepuscular: Occurring daily during the phases of
thermal core through the thermal shell to the heat dis- twilight. → diurnal. (L. crepusculum-twilight) .
sipating surfaces of the body is a thermoeffector of au-
tonomic temperature regulation. Its efficiency in Critical temperature for evaporative heat loss. →
transferring heat is a function of flow rate and the tem- Critical temperature, upper.
perature gradient between core and heat dissipating
surface, and may be modified by the degree of heat ex- Critical temperature for heat production.→ Critical
change by conduction between the vessels to and from temperature, lower.
these surfaces (e.g., due to the counter-current arrange-
ment of arteries and veins). Critical temperature, lower: The ambient tempera-
ture below which the rate of metabolic heat produc-
Convective heat transfer (C): The net rate of heat tion of a resting thermoregulating tachymetabolic
transfer in a moving gas or fluid (i.e., by convection) animal must be increased by shivering and/or nonshiv-
between different parts of an organism, or between an ering thermogenesis in order to maintain thermal bal-
organism and its external environment; it may develop ance. Synonym: critical temperature for heat
and be amplified by thermal gradients (natural convec- production. [°C]
tion) and by forces such as wind, fans, pumps or body
movement (forced convection) usually expressed in Critical temperature, upper: The ambient tempera-
terms of unit area of the total body surface (→ Area, to- ture above which the rate of evaporative heat loss of a
tal body), i.e., as a heat flux. The quantity (C) in the resting thermoregulating animal must be increased
body heat balance equation in which (–C) = heat gain (e.g., by thermal tachypnea or by thermal sweating)
and (+C) = heat loss. [W·m–2], [W], [W·m–3], [W·kg–1] in order to maintain thermal balance. Synonym: criti-
cal temperature for evaporative heat loss. [°C]
Convective heat transfer coefficient.→ Heat trans- Note: Upper critical temperature has also been used
fer coefficient, convective. to define the ambient temperature above which the
capacities of the mechanisms of heat transfer to the
Convective mass transfer. → Mass transfer, convec- environment of a resting animal are exceeded, such that
tive. core temperature is forced to rise and, as a conse-
quence, metabolic energy transformation, i.e., the in-
Convulsions, febrile: Convulsions (epileptic seizures) ternal heat load, is further increased. This use of the
occurring before or during fever episodes. Febrile con- term should be avoided, however, because it does not
vulsions occur mostly in children between the age of 6 accurately describe the upper temperature survival
months and 3 years. They are rare after the age of 8 limit.
years.
Note: Due to its rare occurrence, this term has become Cryogen: An endogenous or exogenous substance that
obsolete in many developed countries. lowers the set-point of body temperature and causes
anapyrexia. Antonym: pyrogen. (Gk. kruos-cold; <
Core temperature. → Temperature, core. gen-become).
Core, thermal: Those inner tissues of the body whose Cryothermy: The thermal status of a supercooled or-
temperatures are not changed in their relationship to ganism (i.e., with the temperature of the body mass be-
each other by circulatory adjustments and changes in low the freezing point of the tissue). (Gk. kruos-cold;
heat dissipation to the environment that affect the ther- therme-heat)
mal shell of the body.
Note: The term is used mainly to describe the deep body Cutaneous evaporative heat loss (CEHL): Rate of
tissues of homeotherms whose temperatures are repre- heat dissipated by evaporation from the skin. [W],
sented by the core temperature. The brain is generally [W·m–2], [W·m–3], [W·kg–1]. Cutaneous evaporative
heat loss (CEHL) plus respiratory evaporative heat of activity) occurring during the day, as distinct from
loss (REHL) constitute → evaporative heat loss. the night. 2. Occurring daily (during each 24-h period).
(L. diurnus adj < dies-day).
Cutaneous water exchange, passive. → Passive cuta- Note: Terms relating to other phases of entrained 24 h
neous water exchange. periodicity, i.e., to definition #1, are: nocturnal (during
the night) and crepuscular (during twilight); nych-
Cutaneous water vapor exchange, passive. → Pas- themeral corresponds to definition #2.
sive cutaneous water vapor exchange.
Dry bulb temperature. → Temperature, dry bulb.
Cytokine: A class of immunoregulatory polypeptides
produced by various cell types, but, at the outset, pri- Dry heat loss. → Heat loss, nonevaporative.
marily by mononuclear phagocytes activated by vari-
ous signals provided by exogenous pyrogens. DuBois area. → Area, DuBois.
Cytokines may act as endogenous mediators of fever
(endogenous pyrogens). The most prominent among Eccritic body temperature. → Preferred body tem-
the fever producing cytokines are interleukins perature.
(IL)-1β and -6, tumor necrosis factor (TNF)-α, and
interferon (IFN)-α. Ectothermic temperature regulator. → Tempera-
ture regulator.
Deep body temperature. → Temperature, core.
Ectothermy: The pattern of temperature regulation
Density (ρ): The ratio of the mass to the volume of a of animals in which body temperature depends mainly
substance. [kg · m–3] on the behaviorally controlled exchange of heat with
the environment. Autonomic thermoeffectors may be
Dew-point temperature. → Temperature, temporarily important in a few ectothermic species
dew-point. (panting in lizards, warming-up of insects). (Gk. ek-
tos-outside; therme-heat) Antonym: endothermy.
Diet-induced thermogenesis. → Thermogenesis,
diet-induced. Effective radiant field. → Radiant flux, effective.
Diffusivity, mass (D): The constant of proportionality Effective radiant flux. → Radiant flux, effective.
relating the rate of diffusion of a gas to the gradient of
its concentration in another gas, e.g., water vapor in air. Effective radiating area. → Area, effective radiat-
·
If the rate of diffusion of mass is m[kg · s–1] in a direc- ing.
tion specified by a coordinate y [m] and if the concen-
tration over a plane at a given value of y is p[kg ·m–3], Effective temperature. → Temperature, effective.
the rate of flow through an area A is given by the Fick-
· Emissivity (ε): The ratio of the total radiant energy
ian equation, m = –DA (∂p/∂y), where D is the mass
diffusivity or diffusion coefficient and ∂p/∂y is the ap- emitted by a body to the energy emitted by a full radi-
propriate concentration gradient. More complex equa- ator at the same temperature.
tions are needed to describe diffusion in two or three
directions simultaneously [m2 · s–1] Emissivity, directional (ε(θ,φ)): The ratio of the ther-
mal radiance (Le,th) of a body in a given direction to
Diffusivity, thermal (α): The thermal conductivity of that of a full radiator (Le,(ε=1)) at the same temperature.
a substance divided by its density (ρ) and specific heat ε(θ,φ) = Le,th(θ,φ)/Le,(ε=1) in which θ and φ are the angular
at constant pressure (cp) ·[m2 · s–1] coordinates defining the given direction.
Direct calorimetry. → Calorimetry, direct. Emissivity, hemispherical (εh): The ratio of the total
radiant energy emitted by an element of a surface into
Directional emissivity. → Emissivity, directional. a hemisphere to the energy by a similar element on the
surface of a full radiator. The element forms the center
Diurnal: 1. Indicates biological processes (e.g., phases of the equatorial plane of the hemisphere, but it is not
necessary to define its radius. components in the body heat balance equation.
Emissivity, spectral (ελ): The ratio of the radiant flux Energy metabolism. → Metabolism, energy.
emitted by an element of surface per unit wavelength
interval to the flux emitted by a full radiator at the Environment. → Total environment.
same temperature and in the same waveband.
Estivation: A state of lethargy (often during the sum-
Emissivity, window (εw): The ratio of the radiant en- mer) with a reduction in body temperature and metab-
ergy emitted by an element of surface between wave- olism demonstrated by some animals that are
length λ1 and λ2 to the flux emitted by a full radiator at temperature regulators when active. The term is usu-
the same temperature and in the same waveband. The ally, but not exclusively, applied to ectothermic tem-
quantity is a special type of spectral emissivity and is perature regulators. (L. aestivus adj <
used because some biological materials have a low aestas-summer)
emissivity in parts of the infrared or visible spectrum.
These parts are known as windows. Eurythermy: The tolerance by organisms of a wide
range of environmental temperatures, or the accommo-
Emittance. → Radiant exitance. dation to substantial changes in the thermal environ-
ment. (Gk. eurus-wide; therme-heat) Antonym:
Endogenous antipyretic. → Antipyretic, endoge- stenothermy.
nous.
Euthermy. → Cenothermy.
Endogenous pyrogen. → Pyrogen, endogenous.
Evaporative heat gain: → evaporative heat transfer
Endothermic temperature regulator. → Tempera- from the ambiance to the body (i.e., gain of heat en-
ture regulator. ergy) due to condensation of vapor on the skin and/or
the surfaces of the respiratory tract, usually expressed
Endothermy: The pattern of thermoregulation in in terms of energy per unit time and unit area of total
which the body temperature depends on a high (ta- body surface. [W], [W · m–2], [W · m–3], [W · kg–1]
chymetabolic) and controlled rate of heat production. Note: evaporative heat transfer most frequently oc-
Behavioural responses are often used by endotherms. curs by vaporization of water from body surfaces and is
(Gk. endo-inside; therme-heat) Antonym: ectothermy. → evaporative heat loss.
Note: The use of endothermy to denote the production
of heat within an organ or organism is etymologically Evaporative heat loss: → evaporative heat transfer
correct but may be found confusing as the same term is from the body to the ambiance (i.e., loss of heat energy)
used in chemistry to mean the uptake of heat during a by evaporation of water from the skin and the surfaces
chemical reaction. of the respiratory tract, usually expressed in terms of
heat flow [W] or heat flux [W·m–2] (energy per unit
Endotoxin: Heat stable compounds that are intrinsic to time and unit area of total body surface) or in [W·kg–1]
the cell walls of Gram-negative bacteria. The term de- or [W·m–3].
rives from the noxious effects on the infected host; in Note: Evaporative heat loss comprises passive com-
addition, endotoxins act as potent exogenous pyrogens ponents, i.e., water vaporizing from respiratory sur-
and stimulators of the acute-phase response, respec- faces at normal respiration and water diffusing through
tively. Endotoxins contain lipopolysaccharides (LPS) the skin and vaporizing at the surface (→ insensible
of high molecular weight. Other cell wall components, perspiration); its thermoregulatory components are
e.g., peptidoglycans, may have partially similar effects. established by autonomic thermoeffectors, like ther-
mal sweating and thermal panting, and behavioral
Energy: Energy may occur either as chemical, electro- thermoeffectors, like saliva spreading, wallowing,
magnetic, or mechanical energy. [Ws = J]. Synonymous and other modes of behavioral surface wetting.
to mechanical energy (force times length, height or dis-
tance) is work, synonymous to thermal energy is heat. Evaporative heat loss coefficient. → Heat loss coef-
Work or heat per unit time (→ work rate or → heat ficient, evaporative.
flow) constitute physically → power, and are used as
Evaporative heat loss, respiratory. → Respiratory named pyrogens. Endogenous pyrogens (e.g., pros-
evaporative heat loss. taglandin E2) are usually the obligatory mediators of fe-
Evaporative heat transfer (E): The rate at which heat ver, but direct actions of exogenous pyrogens are not
energy is transferred by evaporation from or condensa- ruled out. Fever can also result as an idiopathic condi-
tion on the skin and the surfaces of the respiratory tract, tion of the body. It can also be caused by emotional
usually expressed in terms of unit area of total body sur- stress (→ stress fever). The exact mechanisms of fever
face. The quantity (E) in the body heat balance equa- induction are not yet clear.
tion, evaporation and heat loss from the body are Note: Typical natural fever in response to bacterial in-
indicated by (+E), condensation and heat gain to the fection is induced by interleukin-1 (IL-1), IL-6 and
body by (–E). [W] or [W · m–2]. other cytokines which are formed as mediators in the
multifactorial humoral host-defence response termed
Exertional heat stroke: An acute syndrome caused by acute-phase response. Limited febrile rises in core
an excessive rise in body temperature as the result of temperature are apparently harmless, as long as bodily
overloading or failure of the thermoregulatory system functions are not compromised by other disturbances or
during exposure to heat stress while exercising in a stressful conditions. Fever may contribute to the bene-
warm environment. ficial effects of the humoral host-defence responses to
Note: Exertional heat stroke, in contrast to classical infection.
heat stroke, occurs most commonly in healthy young
or middle-aged adults; cutaneous vasodilatation is re- Fever, stress. → Stress fever.
duced and thermal sweating may decrease but is often
still conspicuous. Freezing cold injury (FCI): Tissue damage (frostbite)
resulting when tissue temperature falls below its freez-
Exitance. → Radiant exitance. ing temperature (ca. –2°C). 90% of FCI’s occur in ex-
posed skin and the extremities. The development of
Exogenous pyrogen. → Pyrogen, exogenous. FCI is primarily determined by the ambient tempera-
ture, Antonym: Non-freezing cold injury (NFCI).
Febrile: A term used to describe the state of an organ-
ism during a fever. Antonym: afebrile Full radiator. → Radiator, full.
Fever: A state of elevated core temperature which is Genotypic adaptation. → Adaptation, genotypic.
often, but not necessarily, part of the defensive re-
sponses of organisms (host) to the invasion by live (mi- Globe temperature. → Temperature, globe.
croorganisms) or inanimate matter recognized as
pathogenic or alien by the host. The rise in core tem- GDP-binding protein. → UCP1.
perature is usually designated as due to a change in the
thermocontroller characteristics resulting in an eleva- Gular fluttering: The rapid oscillations of the hyoid
tion of the set-point of body temperature. The higher apparatus and hence of the gular region of some birds as
temperature is actively established and defended by the a characteristic form of thermal panting during expo-
operation of heat-producing and heat-conserving ther- sure to high ambient temperature, by which means air
moeffectors. Time courses and extent of natural fevers is moved across the moist surfaces of the upper respira-
are variable, but an upper temperature limit (41°C in tory tract.
humans) is seldom exceeded, at which level core tem-
perature is maintained for some time and eventually Habituation: Reduction of responses to or perception
reduced when the set-point of temperature regulation of a repeated stimulation.
returns to its normal level. While any rise in body tem-
perature may be due to fever, those rises which are not Heat. → Energy.
accompanied by supportive changes in thermoeffector
activities, are termed hyperthermia. The humoral Heat balance equation. → Body heat balance equa-
mechanisms underlying fever consist of the alteration tion.
of the neural control of body temperature by agents
Heat capacity: The product of the mass of an object refers to convection only and does not include other
and its specific heat. [J ·°C–1] forms of heat transfer.
Heat content, body: The product of the body mass, its Heat loss, evaporative. → Evaporative heat loss.
average specific heat, and the absolute mean body
temperature. [J] Heat loss, insensible. → Evaporative heat loss.
Note: The actual value of this term is seldom calcu- Note: The term insensible heat loss should be avoided
lated. It is used only in the determination of heat stor- in favour of evaporative heat loss, because of possible
age. → Storage of body heat. confusion with insensible water loss which implies
evaporative heat loss but relates to fluid balance.
Heat cramps: Painful spasms of skeletal muscles re-
lated to prolonged heat stress. Heat loss, Newtonian. → Heat loss, dry.
Note: Heat cramps are usually strong painful tetanic
contractions of groups of extremity or abdominal mus- Heat loss, nonevaporative. → Heat loss, dry.
cles. They are related to prolonged heat stress, pro-
found thermal sweating and water but not salt deficit Heat loss, sensible. → Heat loss, dry.
replacement.
Heat production, metabolic. → Metabolic heat pro-
Heat endurance.→ Heat toleranc. duction.
Heat exhaustion: Muscular weakness, fatigue, and Heat production (excess), postprandial: The increase
distress, resulting from prolonged exposure to heat. in metabolic heat production, relative to its postab-
Core temperature is elevated and thermal sweating sorptive resting level, in the hours following food in-
and cutaneous vasodilatation are commonly but not in- take. Its relationship to specific dynamic effect, or
variably reduced. This condition is aggravated by mus- diet-induced thermogenesis is not clear.
cular exertion, dehydration and hyponatremia. Note: While the term postprandial heat production is
Circulatory abnormalities may occur. coming into use, it is recommended to include the spec-
ification excess, because the definition means heat pro-
Heat flow [H]: The amount of heat transferred per duction in excess of resting metabolic rate (RMR).
unit time between parts of a body at different tempera-
tures, or between a body and its environment when at Heat production, total. → Total heat production.
different temperatures. [W]. Physically it constitutes →
power. Heat shock protein (HSP): Intracellular proteins
present in every species, from bacteria to humans.
Heat flow balance equation.→ Body heat balance Among the multiple functions ascribed to these pro-
equation. teins they protect nascent proteins, halt proteins
de-folding etc. Their generation may alter the threshold
Heat flux: Heat flow passing through unit area of a for thermal injury.
given surface. [W · m–2]
Heat shock response. → Thermotolerance.
Heat gain. → Heat transfer.
Heat, specific. → Specific heat.
Heat loss. → Heat transfer.
Heat storage, change in: → Storage of body heat.
Heat loss coefficient, evaporative. → Heat transfer
coefficient, evaporative. Heat stroke: An acute syndrome caused by an exces-
sive rise in core temperature as the result of overload-
Heat loss, dry: The sum of heat flows or heat fluxes ing or failure of the thermoregulatory system during
by radiation, convection, and conduction from a body exposure to heat stress. It is characterized by a large va-
to the environment. [W] or [W · m–2] Synonyms: sen- riety of pathophysiological alterations of bodily func-
sible heat loss; heat loss, Newtonian. tions, including mental disturbances, with a high
Note: In meteorological literature, sensible heat loss incidence of permanent or fatal damage.
Note: Classical heat stroke, in contrast to exertional combination of these processes) and for evaporation
heat stroke, occurs under sedentary conditions, most (see below).
commonly in very old or very young people; cutaneous
vasodilatation is greatly reduced and thermal sweating Heat transfer coefficient, combined nonevaporative
is commonly absent. (hcomb): The ratio of total heat transfer per unit area
by radiation, convection, and conduction to the temper-
Heat syncope: Collapse, usually with loss of con- ature difference between the surface and operative
sciousness, during exposure to heat. The symptoms are temperature of the environment.
similar to those of the vasovagal syndrome (fainting).
hcomb = hr + hc + hk [W· m–2 · °C–1]
Heat tolerance: The ability to tolerate high ambient
temperatures. This term comprises a variety of physi- Heat transfer coefficient, conductive (hk): The net
ological characteristics. Homeotherms (tachymeta- heat transfer by conduction per unit area between a
bolic species) are often characterized as heat tolerant, surface and a solid or stationary fluidic medium in con-
if they remain comfortable or are able to balance heat tact with the surface per unit temperature difference
production and heat loss at particularly high ambient (∆T) between the surface and the substance with which
temperatures. Homeothermic species are also char- it is in contact.
acterized as heat tolerant, if they are capable to main-
tain normal functions, or to survive, at body hk = K· ∆T–1 [W·m–2 ·°C–1]
temperatures exceeding their normal range; this may
involve the facility of selective brain cooling. – Poiki- Heat transfer coefficient, convective (hc): The net
lotherms (bradymetabolic species) may also be spec- heat transfer per unit area between a surface and a
ified as heat tolerant, if they are able to survive very moving fluidic medium per unit temperature difference
high body temperatures. All forms of heat tolerance (∆T) between the surface and the medium.
may be subject to adaptation or acclimat(izat)ion.
Synonym: heat endurance. hc = C· ∆T–1 [W· m–2· °C–1]
Note: In the OED the words tolerance and endurance
are almost synonymous. From a physiological point of Heat transfer coefficient, evaporative (he): The net
view a distinction is sometimes made between heat tol- heat transfer per unit vapor pressure gradient caused
erance and heat endurance, the former term is some- by the evaporation of water from a unit area of wet sur-
times used in a more general sense and includes all face or by the condensation of water vapor on a unit
forms of tolerance, while the latter is sometimes used in area of body surface. The driving force is the vapor
a more specific sense as in tolerance with time during a pressure gradient from Pws (on the surface) to Pwa (of
sustained stress, e.g., exercise endurance. the ambient gas). Thus
Heat transfer: Process of heat flow [W], i.e., the rate he = E(Pws – Pwa)–1· [W· m–2· Pa–1] or [W· m–2 ·Torr–1].
at which heat energy is transferred from one part of an
organism to another especially between body core and In terms of the latent heat of vaporization (λ or LHV)
shell. In this case the heat transfer is a combination of of water, the gas constant for water vapor (Rw), the
conduction and convection. A more frequently investi- mean temperature (T) of the medium in °K and the
gated topic of thermal physiology is heat transfer be- mass transfer coefficient (hD)
tween an organism and its environment, by
conduction, convection, radiation, evaporation, or a he = hD·λ·Rw–1·T–1
combination of these. Net transfer from the organism to
the environment is heat loss, and from the ambiance Note: In most physiological applications the vapor
into the organism is heat gain. → Body heat balance pressure gradient instead of the concentration gradient
equation. can be considered as the driving potential for the evap-
orative process since the temperature difference be-
Heat transfer coefficient: A parameter (h) [W · m–2 tween the evaporating surface and the ambient air is
·°C–1] which determines the amount of heat flux (H) small in relation to the average temperature of the sur-
due to a temperature difference (∆T): H = h· ∆T. H may face-water vapor medium.
be defined for conduction, convection, radiation (or the
Heat transfer coefficient, radiative: The net rate of perature and metabolism of some animals that are ho-
heat transfer per unit area by the exchange of thermal meothermic temperature regulators when active. (L.
radiation between two surfaces, per unit temperature hibernare- to pass the winter)
difference between the surfaces. [W · m–2 · °C–1] Note: Body temperature is regulated in the state of hi-
bernation, however, with set-point differing from that
Heat transfer coefficient, radiative (linear) (hr): Ac- in the state of cenothermy (euthermy).
cording to the Stefan-Boltzmann Law, the exchange
of radiation between two black surfaces at temperatures Hidromeiosis: The swelling of keratinized layers of
T1 and T2 [°K] is proportional to σT4, where σ is the the skin, due to prolonged exposure to water or sweat,
Stefan-Boltzmann constant. When the ratio (T1 – T2)/T which blocks sweat ducts and reduces sweating rate.
is small, where T = (T1 + T2)/2,σ(T14 – T24) ≈ 4σT3 · (Gk. hidros-sweat; meiosis-decrease)
(T1 + T2) and the term 4σT3 can be treated as a linear
heat transfer coefficient. [W · m–2 · °C–1] Homeostasis: General term characterizing the relative
constancy of physico-chemical properties of the inter-
Heliothermy: The regulation of the body temperature nal environment of an organism as being maintained by
of an ectothermic animal by behavioral adjustments of regulation. (Gk. homoio-similar, resembling; sta-
its exposure to solar radiation. (Gk. helios-the sun; sis-standing)
therme-heat)
Homeothermy: The pattern of temperature regula-
Hemihidrotic response: Suppression of thermal tion in a tachymetabolic species in which the cyclic
sweating, due to the skin pressure effect, if one half of variation in core temperature, either nychthemerally
the body is affected. or seasonally, is maintained within arbitrarily defined
limits despite much larger variations in ambient tem-
Hemispherical emissivity. → Emissivity, hemispher- perature, i.e., homeotherms regulate their body tem-
ical. perature within a narrow range. (Gk. homoio-similar,
resembling; therme-heat) Synonym: homoiothermy.
Heterothermy: The pattern of temperature regula- Note: A number of homeothermic species may tempo-
tion in a tachymetabolic species in which the variation rarily enter states in which their set-range of body tem-
in core temperature, either nychthemerally or sea- perature regulation is not definable or much lower
sonally, exceeds that which defines homeothermy. than during their normal state of homeothermy (→
(Gk. hetero-different; therme-heat) heterothermy, hibernation, torpor).
Note: This is an arbitrary term in two senses: i) because
hetero = a difference from (i.e., a difference between Homoiothermy. → Homeothermy.
two) rather than a wide range, and ii) because the dis-
tinction that is being made between thermostable and Humidity, absolute (γ): Mass of water vapor in air per
rather less thermally stable species depends on an arbi- unit volume of air/water vapor mixture. [kg · m–3]
trary division. →: 1. poikilothermy (diversified tem-
peratures) although etymologically satisfactory has Humidity, relative (φ): The ratio of the mol fraction of
already been given a more restricted meaning. 2. steno- water vapor present in a volume of air to the mol frac-
thermy (narrow temperature range) and eurythermy tion present in saturated air, both at the same tempera-
(wide temperature range) would have suited the cir- ture and pressure; in thermal physiology, the ratio of the
cumstance well, but these terms are already in use to saturated vapor pressure at the dew point tempera-
describe animals that are able to adjust to narrow ranges ture (Ps, dp) of the enclosure to the saturated vapor
and wide ranges respectively, of ambient tempera- pressure at its dry bulb temperature (Ps, db) [ND].
ture. When the relative humidity is expressed as a percent-
age, the symbol is RH.
Heterothermy, local: The pattern of temperature in
those parts of the body which comprises the thermal Hyperpnea, thermal. → Thermal hyperpnea.
shell of homeotherms.
Hyperthermia: The condition of a temperature regu-
Hibernation: The state of winter (sometimes late fall lator when core temperature is above its range speci-
or early spring) lethargy with a reduction in body tem- fied for the normal active state of the species. (Gk.
Interthreshold zone. → Thermoeffector threshold Mass transfer, convective: The transport by convec-
zone. tion of one component of a nonreactive mixture (usu-
ally air-water) across an interface caused by a
Irradiance (E): The radiant flow incident on or pass- concentration gradient often accompanied by a trans-
ing through unit area of a surface. [W · m–2] formation of phase and by a simultaneous transfer of
heat.
Katathermometer: An instrument which measures the
heat flow from a surface to a cooler environment. ·
Mass transfer rate (m): The rate of transfer of mass.
–1
[kg · s ]
Latent heat of fusion: The quantity of heat absorbed,
or released, in the reversible change of state, melting or Maximum metabolic rate. → Metabolic rate, maxi-
freezing, of unit mass of solid or liquid, without change mum.
of temperature. [J · g–1]
Note: The melting of 1 g of ice to water at 0°C absorbs Maximum oxygen consumption. → Oxygen con-
333.15 J. sumption, maximum.
Latent heat of vaporization (λ or LHV): The quantity Mean body temperature. → Temperature, mean
of heat absorbed (or released) by a volatile substance body.
(fluid) per unit mass in the process of its reversible
change of state by evaporation (or condensation) under Mean radiant temperature. → Temperature, mean
isobaric and isothermal equilibrium conditions. [J · g–1] radiant.
Note: For water: λ = 2490.9 – 2.34 T, with T = temper-
ature of water in °C, e.g., l g of water, when changing Mean skin temperature. → Temperature, mean
from fluid to vapor at T = 34°C, absorbs 2411.3 J. skin.
Least observed metabolic rate. → Metabolic rate, Menopausal hot flush (flash): An abrupt heat dissipa-
least observed. tion response, provoked or unprovoked, occurring in
hypoestrogenic women following the decline of ova-
Lower critical temperature. → Critical tempera- rian function. The response typically lasts about 5 min-
ture, lower. utes and consists of peripheral vasodilation, sweating
and reports of intense warmth.
Lower temperature survival limit. → Temperature
survival limit, lower. Met: An assigned unit of measurement to designate
"sitting-resting" metabolic rate of man.
Malignant hyperthermia. → Hyperthermia, malig-
nant. 1 met = 58.15 W · m–2 = 50 kcal · h–1 · m–2
W3/4 as the metabolic body size permits comparisons Metabolic rate (M or MR): = metabolic energy
to be made between the metabolic levels of different transformation. [W], [W·m–2], [W·m–3], [W·kg–l],
animals. When W2/3 is used, this implies proportional- [W·kg–3/4] The rate of transformation of chemical en-
ity of metabolic body size to the animal's surface area ergy into heat and mechanical work by aerobic and
(→ surface rule). The relation between metabolic rate anaerobic metabolic activities within an organism, usu-
(M or MR) and body size (M = aW3/4) is a particular ally expressed in terms of unit area of the total body
case of the general allometric equation (y = axb) which surface, i.e., as a heat flux. The quantity M in the body
says that plotting a biological variable, y, against an- heat balance equation.
other biological variable, x, after logarithmic transfor- Note: Metabolic energy transformation may not all
mation (i.e., log y = log a + b · log x) will result in a result from aerobic metabolic activities and may there-
straight line with slope b. Ref: von Bertalanffy, L., Hel- fore exceed that indicated by the rate of oxygen con-
goländer Wiss. Meeresuntersuch. 9: 5, 1964. sumption. Part of the metabolic energy
transformation may be used to do work on an external
Metabolic energy production. → Metabolic energy system, and therefore the rate of heat production may
transformation. be less than the metabolic energy transformation.
Terms in the body heat balance equation are often ex-
Metabolic energy transformation (M). → Metabolic pressed as quantities of energy per unit surface area
rate. and per unit time [W · m–2], because heat exchange is a
function of area. Metabolic rate (M or MR) is given as
Metabolic heat production (H): Rate of transforma- the total energy production in the organism in unit time
tion of chemical energy into heat in an organism, usu- [W] or often as the energy production per unit mass of
ally expressed in terms of unit area of the total body tissue in unit time [W · kg–1]. For comparison of meta-
surface (→ total body area). H is equal to M – (W) in bolic rates of animals of different body sizes, meta-
the body heat balance equation. [W · m–2], [W · kg–1] bolic rate (M or MR) is usually related to (body
or [W] mass)3/4 (→ metabolic body size).
Note: During positive work rate (+W) or in the ab-
sence of work (W = 0) metabolic heat production (H = Metabolic rate, basal (BMR): metabolic energy
M – (+W)), i.e. positive work rate subtracted from the transformation calculated from measurements of heat
rate of metabolic energy transformation (=meta- production or oxygen consumption in an organism in a
bolic rate), equals total heat production. When work rested, awake, fasting* sufficiently long to be in
is being done on the body by an external source (nega- post-absorptive state, and thermoneutral zone (a par-
tive work), total heat production is the sum of meta- ticular case of standard metabolic rate (SMR)).[W],
bolic rate (M or MR) (which is equal to metabolic heat [W · m–2], [W · m–3], [W · kg–1], [W · kg–3/4]
production in this condition) and the heat liberated Note: In these conditions, when the amount of work be-
within the body due to negative work (M – (– W)). ing done on an external system is negligible, the rate of
heat production is equal to the rate of metabolism
Metabolic level: The metabolic heat production (metabolic energy production).* The period of fast-
measured under standard conditions (→ metabolic ing needs to be specified as this may be for days in large
rate, standard (SMR)) during a 24-h period divided by animals, and for much shorter periods for very small
the metabolic body size. [kJ · kg–3/4· (24 h)–1]* mammals and birds.
Note: *1 kJ . kg–3/4 · (24 h)–l = 0.2388 kcal · kg–3/4 ·(24
h)–l. Metabolic levels are approximately constant Metabolic rate, lowest observed (LOMR): The lowest
within phylogenetic groups but may vary between observed rate of metabolism during specified periods
groups. For example, mammalian and avian species of minimum activity. [W], [W · m–2], [W · kg–1], [W ·
(tachymetabolism) have higher metabolic levels than kg–3/4] → metabolic rate, minimum observed
other species (bradymetabolism), while the metabolic (MOMR). The rationale and objective of lowest ob-
levels of birds are apparently higher than those of mam- served metabolic rate (LOMR) and minimum ob-
mals, and those of prototherian (monotreme) and met- served metabolic rate (MOMR) are identical: to
atherian (marsupial) mammals are lower than those of measure the metabolic rates of small and wild animals
eutherian (placental) mammals. during periods of minimal activity as the nearest that
(Poczopko, 1971). Ref: Poczopko, P., Acta Theriolog- can be made to a measurement of a standard meta-
ica 16: 1, 1971. bolic rate (SMR).
Note: There are small but significant differences in [W], [W· m–2], [W · m–3], [W · kg–1], [W · kg –3/4]
technique: Minimum observed metabolic rate Note: A particular case of standard metabolic rate
(MOMR) is the average metabolic rate (M or MR) (SMR) used when the subject cannot be brought to a
during periods of minimum activity; lowest observed fasting condition, e.g., ruminant animals. The period of
metabolic rate (LOMR) is the lowest recorded meta- food deprivation should be stated.
bolic rate (M or MR) during periods of minimum ac-
tivity. Such brief low values may be influenced by Metabolic rate, standard (SMR): metabolic energy
physical characteristics of the system of measurement. transformation calculated from measurements of heat
Note: lowest observed metabolic rate (LOMR) may production or oxygen consumption in an organism un-
be lower than basal metabolic rate (BMR) (e.g., dur- der specified standard conditions*. [W], [W · m–2], [W
ing some stages of sleep). · m–3], [W · kg–1], [W · kg–3/4]
Note: The conditions are usually such that the amount
Metabolic rate, maximum (MMR): The highest met- of work being done on an external system is negligible.
abolic rate (M or MR) during a specified period of The rate of heat production is then an acceptable index
work compatible with sustained aerobic metabolism of the rate of metabolism. * The specified standard con-
(i.e., when there is no progressive accumulation of lac- ditions are usually that the organism is rested (or as near
tic acid in the blood). [W], [W · m–2], [W · m–3], [W · to rested as is possible), fasting (if possible), awake,
kg–1],[W · kg–3/4] and in a thermoneutral environment. The extent to
Note: There may be some confusion between maxi- which standard conditions can be achieved varies with
mum and peak metabolic rates (PMR). Both terms in- species. → Metabolic rate, minimum observed
dicate maximum rates. The distinction is in usage: (MOMR).
maximum relates to work, peak relates to cold expo-
sure. The terms should also be distinguished from max- Metabolic rate, summit. → Metabolic rate, peak.
imum oxygen consumption.
Metabolic scope. → Scope, metabolic.
Metabolic rate, minimum observed (MOMR): Aver-
aged metabolic rate during specified periods of mini- Metabolically effective body mass: → metabolic
mum activity. [W], [W · m–2], [W · m–3], [W · kg–l], [W body size (preferred synonym).
· kg–3/4] Note: The term metabolically effective body mass
Note: The metabolic rate (M or MR) of small animals may be wrongly understood to be that part of the body
in particular, but also of larger wild animals, cannot be mass which is metabolically active in contrast to a part
measured under basal or other standard conditions (→ that is metabolically inert. Its use should be avoided.
metabolic rate, basal (BMR), and metabolic rate,
standard (SMR)). A practical solution to the problem Metabolism. → Metabolism, energy. (Gk. metab-
is to measure metabolic rate (M or MR) continuously ole-change)
and accept the average metabolic rate (M or MR) dur- Note: Metabolism is a general term which relates to
ing periods of minimum activity as the best possible es- chemical and physical changes occurring in living or-
timate of standard metabolic rate (SMR). ganisms. In thermal physiology metabolism invariably
relates to the transformation of chemical energy into
Metabolic rate, peak (PMR): The highest metabolic work and heat. In other divisions of physiology the
rate (M or MR) that can be induced in a resting animal term is used in relation to other changes in state of
by any cold environment. [W], [W · m–2], [W · m–3], chemical energy (e.g., from storage compounds to
[W · kg–l], [W · kg–3/4] Synonym: summit metabolic readily available compounds, including the energy
rate. → Metabolic rate, maximum (MMR). transferring polyphosphates, and vice versa), or to turn-
Note: Although summit metabolic rate (SMR) is now an over of material between different states (e.g., calcium
accepted term, peak metabolic rate (PMR) is prefera- salts from ionized to solid, and vice versa).
ble because the abbreviation SMR is indistinguishable
from that for standard metabolic rate (SMR). Metabolism, anaerobic: Transformation of matter and
energy without uptake of oxygen.
Metabolic rate, resting (RMR): The metabolic rate
(M or MR) of an animal that is resting in a thermoneu- Metabolism, energy: The sum of the chemical
tral environment but not in the postabsorptive state. changes in living matter in which energy is trans-
Microwave: Part of the electromagnetic spectrum with Operative temperature. → Temperature, operative.
wavelength from 1 to 100 mm. Microwaves are ab-
sorbed by metal and water and thus permit deep heating Optimal body temperature range: The range of body
of aqueous or metallic bodies exposed to them (→ ra- temperatures in which a species carries out its normal
diant energy). daily activity.
Note: This “ecological optimum” integrates internal
Minimum observed metabolic rate. → Metabolic and external forces acting on a species; it applies
rate, minimum observed. mostly to ectotherms.
·
Natural convection. → Convective heat transfer. Oxygen consumption, maximum (VO2max): The
maximum rate at which an organism can take up oxy-
Negative work. → Work rate, negative. gen. [ml · s–1], conventionally [ml · min–1], [l · min–1]
Note: Determination of this parameter requires very
Neuroleptic malignant syndrome: The development high motivation of the subject and can probably be
of severe muscle rigidity, elevated core temperature, done only on humans. Criteria used to show that a hu-
and a disturbance of mental status associated with the ·
man subject has reached the VO2max although not as yet
use of neuroleptic medication. agreed upon, include an indication of no further in-
crease in oxygen uptake during further increase in work
Nocturnal: Occurring during the nighttime, as distinct load.
from daytime. (L. nocturnus adj < nox-night) Antonym:
diurnal. P 32000 polypeptide. → UCP1
Nonevaporative heat loss. → Heat loss, nonevapora- Panting, thermal. → Thermal panting.
tive.
Partitional calorimetry. → Calorimetry, partitional.
Non-freezing cold injury (NFCI): Tissue damage re-
sulting from prolonged exposure to ambient tempera- Passive cutaneous water exchange: The passage
tures between 15°C down to –0.5°C. In NFCI the through the skin in either direction of water down an os-
normal thermoregulatory responses of the injured body motic gradient per unit area in unit time. [kg . m–2 · s–1],
are disturbed. All types of tissue may be damaged by also [mg · m–2 · s–1]
sustained cooling, but the most serious effects occur in Note: Passive cutaneous water exchange occurs only
the nerves and blood vessels. Antonym: Freezing cold when the skin is covered with water or an aqueous so-
injury (FCI). lution.
Nongenetic adaptation. → Adaptation, phenotypic. Passive cutaneous water vapor exchange: The pas-
sage through the skin in either direction of water vapor
Nonshivering thermogenesis. → Thermogenesis, down a water vapor pressure gradient per unit area in
nonshivering. unit time. [kg · m–2 · s–1], also [mg · m–2 · s–1]. Syn-
onym: insensible perspiration.
Nonthermal sweating. → Sweating, nonthermal.
Passive temperature lability. → Temperature labil-
Normothermy. → Cenothermy. ity, passive.
Nychthemeral: Relating to an exact period of 24 h. Peak metabolic rate. → Metabolic rate, peak.
(Gk. nux-night; hemera-day).
Phenotypic adaptation. → Adaptation, phenotypic.
Nychthemeron: A period of 24 h, consisting of a day
and a night (SOED). Physical temperature regulation. → Temperature
regulation, physical.
Obligatory nonshivering thermogenesis. → Ther-
orological tables and formulas (Ref: Chambers, A.B., A Temperature and Life, Berlin-Heidelberg-New York,
psychrometric chart for physiological research, J. Appl. Springer, 1973, p. 17 ff.).
Physiol. 29: 406–412, 1970). The water vapor pressure
in an enclosure is equal to the saturated vapor pres- Radiance (Le): Radiance at a surface element (dA) of
sure at its dew-point temperature (Ps,Tdp) or to the a source or receiver is the radiant intensity (dI) from
product of the relative humidity (N) and the saturated direction θ divided by the orthogonal projection of this
vapor pressure at its dry bulb temperature (NPs,Tdp). surface element (dA ·cosθ) on a plane perpendicular to
the direction θ. θ is the angle between the normal to the
Projected area. → Area, projected. element (dA) of the source or receiver and the direction
of the observation. [W · sr–1 · m–2]
Pyrexia: The condition of being febrile.
Note: Pyrogen: The generic term for any substance Radiance thermal (Le,th): radiance due to thermal ra-
whether exogenous or endogenous that causes fever diation. [W · sr–1 · m–2]
when introduced into or released in the body. (Gk.
pur-fire; <gen-become). Radiant absorptance, total (α): The ratio of total ra-
diant flux absorbed by a body to the total incident flux.
Pyrogen, bacterial: Any pyrogen derived from bacte-
ria. Radiant emittance. → Radiant exitance.
Note: All endotoxins are bacterial pyrogens.
Radiant energy (Q): Energy travelling in the form of
Pyrogens, endogenous: Heat labile proteins, and lipids electromagnetic waves. [J]
that cause fever. Their production is often stimulated Note: This term should be distinguished from the radi-
by exogenous pyrogens (microorganisms and their ant heat exchange (R) of the environment with the
products, e.g., antigens), but also by injury, trauma and body. That part of the electromagnetic spectrum of sig-
stress. The best identified endogenous pyrogen is in- nificance in thermal physiology is divided for conve-
terleukin-1β (IL-1β). It is thought that this cytokine nience into the wavebands:
induces fever via the production of another cytokine,
IL-6. Other endogenous pyrogens include inter-
feron-α and tumor necrosis factor-α. The final com- Ultraviolet 0.25–0.38 µm
mon pathway for cytokine-mediated fever is generally Visible 0.38–0.78 µm
thought to be the production of prostaglandins of the E
series in or near the anterior hypothalamus. Infrared 0.78–100 µm
Radiant exitance, self (Me,s): The radiant flux emit- Radiant intensity, spectral (Iλ): The radiant inten-
ted by a surface. The flux considered does not include sity per unit wave length interval. [W · sr–1 · nm–1]
reflected or transmitted flux. [W · m–2]
Radiation reflectance. → Reflectance, radiation.
Radiant exitance, thermal (Me,th): The radiant flux
emitted as thermal radiation by a surface. Radiation shape factor (Fij): A dimensionless quan-
Note: In the case of a full radiator, the radiance (Le) is tity expressing the fraction of the diffuse energy emit-
uniform in all directions. In consequence, when the ted by a surface (or a source), denoted by the subscript
solid angle is measured in steradians, the radiant exi- i, that is received by another surface, denoted by the
tance has the numerical value Me = 4πLe. subscript j, visible by it and in known geometric rela-
tion with it. Synonym: radiation view factor.
Radiant field, effective. → Radiant flux, effective.
Radiation transmittance. → Transmittance, radia-
Radiant flow (φ): The amount of radiant energy per tion.
unit of time. [W]
Radiation view factor. → Radiation shape factor.
Radiant flow spectral (φλ): The radiant flow per unit
wavelength interval at wavelength λ. [W · nm–1] Radiative heat transfer coefficient. → Heat transfer
coefficient, radiative.
Radiant fIux (dφ/dA): The radiant flow per unit area.
[W · m–2] Radiative (linear) heat transfer coefficient. → Heat
Note: The radiant flux emitted by a surface is the ther- transfer coefficient, radiative (linear).
mal radiant exitance. The radiant flux incident on or
passing through an area is the irradiance. Radiator: An emitter of radiant energy.
Radiant flux, effective (Hr): The net radiant flux ex- Radiator, full: A radiator of uniform surface temper-
changed with all enclosing surfaces and with any in- ature whose radiant exitance in all parts of the spec-
tense directional sources and sinks for exchange of trum is the maximum obtainable. The emissivity of a
radiant heat by a human or human-shaped object whose full radiator is unity for all wavelengths. Synonym:
surface temperature is hypothetically at ambient air blackbody.
temperature. [W · m–2]
Note: If the same object, with surface temperature at Radiator, graybody: A radiator whose spectral
ambient air temperature, is assumed to exchange ra- emissivity is less than unity, at least in the waveband
diant heat at a given effective radiant flux with a hy- for thermal radiation (3–30 µm), but is the same at all
pothetical black surface enclosure of corresponding wavelengths.
wall temperature, then this condition defines the effec-
tive radiant field transferring the amount of heat per Radiator, selective: A radiator with a spectral emis-
unit time and per unit surface of the object which is sivity less than unity which varies with wavelength.
quantified by the effective radiant flux. In this sense, Note: Human and animal skins have high emissivities
effective radiant field and effective radiant flux are in the waveband 3–30 µm but not between 0.7 and 3
synonymous terms. µm.
Radiant heat exchange (R): The net rate of heat ex- Reflectance, radiation (p): The ratio of the radiant
change by radiation between an organism and its envi- flux reflected by a surface or medium to the incident
ronment, usually expressed in terms of unit area of the flux.
total body surface, i.e., as a heat flux. The quantity (R) Note: Measured values of reflectance depend upon the
in the body heat balance equation where (–R) = heat angles of incidence and view and the spectral character
gain and (+R) = heat loss. [W] or [W · m–2] of the incident flux; these factors should be specified.
Radiant intensity (I): The radiant flow proceeding Regulation: The processes by which a biological sys-
from a source per unit solid angle in the direction con- tem stabilizes variables, generally by information loops
sidered. [W · sr–1] (negative feedback control). The regulated (controlled)
physiological variables can usually be described as a Selective brain cooling: Lowering of brain tempera-
function of physical variables. In the case of changes of ture, either locally or as a whole, below aortic (arterial
a regulated variable due to external or internal interfer- blood) temperature. Cool venous blood returning from
ence, the processes of regulation will alter certain cephalic heat dissipating surfaces acts as a heat sink, ei-
bodily functions, the effectors in the regulatory process, ther for brain tissue directly or for arterial blood supply-
which act to minimize the deviations of the regulated ing brain tissue. Special vascular arrangements, e.g.,
variable from its set-point. the ophthalmic or carotid retia mirabilia, support arte-
Note: In thermal physiology, core temperature is often rio-venous heat exchange underlying selective brain
considered as the regulated variable in homeothermic cooling. Its existence in many species has been proven,
species, because its changes are most effective in acti- but the situation in humans is still debated.
vating the thermoeffectors of heat production, heat
conservation, or heat loss in such a way that deviations Sensible heat loss. → Heat loss, dry.
of core temperature are minimized. Mean body tem-
perature, if representing the weighted contributions of Set-point: The value of a regulated variable which a
core and shell temperatures to the entire thermal infor- healthy organism tends to stabilize by the processes of
mation, may be a better estimate of the regulated vari- regulation. It is found in the condition when the effec-
able. tor activities tending to counteract alterations of the
regulated variable are minimal. With thermal load
Relative humidity. → Humidity, relative. present, the regulated variable, due to proportional con-
trol, does not stabilize at, but near the set-point. In hy-
Resistance, thermal. → Thermal resistance. per- or hypothermia, body temperature deviates
substantially from the set-point, mostly because of in-
Respiratory evaporative heat loss (REHL): Rate of sufficient effector capacity and thereby lack of stabili-
heat dissipated by exhalation of air saturated with water zation. In biological systems, the apparent set-points of
vapor, [W], [W·m–2], [W·m–3], [W·kg–1]. many regulated variables may change with time, either
Note: Respiratory evaporative heat loss (REHL) periodically or temporarily (e.g., nychthemerally, an-
comprises a passive (obligatory) fraction associated nually, with the ovulatory cycle, or in connection with
with normal breathing and, in many animals, a regula- other physiological events). In temperature regula-
tory fraction which depends on the rate at which the wa- tion the set-point may change temporarily, due to inter-
ter vapor dissipating respiratory surfaces are ventilated ference with the regulations of nonthermal variables
(e.g., during thermal tachypnea), and on surface tem- (e.g., in states of dehydration and starvation, etc.), or
perature depending on circulatory convection (→ due to pathological, nonthermal influences (e.g., in fe-
convection, circulatory) transferring heat to these sur- ver or anapyrexia). Also, the processes of accli-
faces. Net dissipation of evaporated water and, conse- mat(izat)ion and adaptation may change the set-point
quently, of heat with the exhaled air is possible only as of temperature regulation. Hibernation is a special
long as the inhaled air is not saturated with water vapor condition in which the set-point differs distinctly from
at the temperature of the moist respiratory surfaces. that existing in the same animal in the state of ceno-
thermy. The change of set-point in these processes is
Resting metabolic rate. → Metabolic rate, resting. thought to be due to changes in the thermal controller
characteristics, particularly changes of thresholds (→
Saliva spreading: The spreading of saliva on the body thermoeffector threshold temperature) and/or
surface, often a deliberate (behavioral) thermoeffector changes of thermoeffector gain, which obviously have
action to cool the surface by evaporation. Sometimes neuronal correlates, too.
inaccurately termed grooming. Note: The term set-point has evoked much confusion,
as it has been used for different phenomena: (1) for
Saturated vapor pressure. →Pressure, water vapor. steady state of body temperatures (which can be very
different in the thermoregulatory system depending on
Scope, metabolic: Metabolic rate (M or MR) at max- environmental and internal conditions). (2) for a central
imum oxygen consumption divided by least observed reference signal (which obviously does not exist explic-
metabolic rate (LOMR). itly in the thermoregulatory system). (3) for thermoef-
fector threshold zone of (mean) body temperature,
Second phase panting. → Thermal hyperpnea. which is correct only in the non-acclimated and afe-
Skin pressure effect on sweating: The inhibition of Steady state: The state of body heat balance in which
sweating in the dermatomal area of skin when mechan- there is no positive or negative heat storage when heat
ical pressure is locally applied (→ hemihidrotic re- gain and heat loss between the body and the environ-
sponse). ment are equivalent. The body temperatures reached in
a steady state depend on the extent of external or inter-
Skin wettedness. → Wettedness, skin. nal thermal load. If the thermoeffector capacities are
exceeded due to excessive load or pathological pro-
Solar radiation area. → Area, solar radiation. cesses, a steady state will not be reached, leading to
Specific dynamic action. → Specific dynamic effect. hypo- or hyperthermia.
Specific dynamic effect: Temporary increase in meta- Stefan-Boltzmann law: The thermal radiant exi-
bolic energy transformation following food intake. tance of a full radiator is proportional to the fourth
power of its absolute temperature, Me,th = σT4. The cur- Sweating, thermal: A response of the sweat glands to
rently recommended value of the Stefan-Boltzmann a thermal stimulus. Rate [mg · m–2 · s–1]
constant (σ) is 5.6696 × 10–8 [W · m–2 · K–4]. Note: 1 mg · m–2 · s–1 = 3.6 g · m–2 · h–1.
Stenothermy: Descriptive of organisms, which occur Sweating topography, thermal: The sequence of the
naturally in a narrow range of environmental tempera- onset of thermal sweating and the differences in sweat
tures and which, singly or collectively, are intolerant of rate observed between different skin regions.
or accommodate ineffectually to wide changes in their
thermal environment. (Gk. stenos-narrow; therme- Tachymetabolism: The high level of basal metabo-
heat) Antonym: eurythermy. lism of birds and mammals relative to those of reptiles
and other nonavian and nonmammalian animals of the
Storage of body heat (S): The rate of increase (+) or same body mass and at the same tissue temperatures.
decrease (–) in the heat content [Ws = J] of the body (Gk. takhus-fast; metabole-change) Synonym:
caused by an imbalance between heat production (met- warm-blooded. Antonym: bradymetabolism,
abolic heat transformation) and heat loss, usually ex- cold-blooded.
pressed in terms of unit area of total body surface (→ Note: This relatively high level of basal metabolism in
Area, total body). The quantity S in the body heat bal- mammals and birds is a precondition for the relative
ance equation. [W], [W · m–2], [W · m–3] or [W · kg–1]: stability of core temperature during exposure to cold
(warm-blooded) and of endothermic homeothermy
Stress fever: A rise in body temperature due to expo- and heterothermy.
sure of a person or an animal to a psychological stres-
sor. Synonym: Stress hyperthermia. Tachypnea, thermal. → Thermal tachypnea.
Stress hyperthermia. → Stress fever. Temperature: A measure of the mean kinetic energy of
the molecules in a volume.
Sunstroke: An acute and dangerous reaction to heat
exposure caused by a failure of the heat regulating Temperature, ambient (Ta): The average temperature
mechanisms of the body. It is characterized by high of a gaseous or liquid environment (usually air or wa-
core temperature, usually above 40.6°C, cessation of ter) surrounding a body, as measured outside the ther-
thermal sweating; headache, numbness, tingling and mal and hydrodynamic boundary layers that overlay the
confusion prior to sudden delirium or coma, fast pulse, body. [°C] Synonym: temperature, dry bulb (in a gas-
rapid respiratory rate and elevated blood pressure. eous environment).
Surface area. → Area, total body, and area DuBois Temperature coefficient: The ratio between the
(for humans only). change in any temperature dependant activity and the
defined temperature range within which this change oc-
Surface rule: A statement that the basal metabolic curs.
rate (BMR) is proportional to the 2/3 power of body
mass. Temperature conformer: An organism, the core tem-
Note: The rule is based on the proposition that basal perature of which varies as a proportional function of
metabolic rate (BMR) is related to surface area and ambient temperature; an animal without effective
that surface area varies with the 2/3 power of body temperature regulation by autonomic or behavioral
mass. However, this is not experimentally verifiable, means. Synonym: poikilothermy. Antonym: tempera-
for when basal metabolic rate (BMR) is expressed per ture regulator.
2/3 power of body mass it increases systematically with
body size (Kleiber, 1947). Basal metabolic rate Temperature conformity: The thermal relation be-
(BMR) is more nearly proportional to the 3/4 power of tween the environment and an organism, the core tem-
body mass (→ metabolic body size). Ref: Kleiber, M., perature of which varies as a proportional function of
Physiol. Rev. 27: 411, 1947. ambient temperature, i.e., an absence of effective
temperature regulation by autonomic or behavioral
Sweating, nonthermal: A response of the sweat means.
glands to a nonthermal stimulus.
Thermal indifference, zone of: The range of ambient temperature regulation; 2. Any activation of thermo-
temperatures, associated with specified water vapor effector activities in response to thermal stress that
pressure, air velocity, and radiant exchange, within cause sustained changes in the state of other, nonther-
which 80% of active people do not complain of the mal, regulatory systems.
thermal environment. [°C] Cf. thermoneutral zone.
Thermal stress: Any change in the thermal relation be-
Thermal inertia for radiant heat ( √kpc ): One of the tween a temperature regulator and its environment
properties of a material which determines the rate of in- which, if uncompensated by temperature regulation,
crease of surface temperature during an exposure to ir- would result in hyper- or hypothermia.
radiance (E). For nonpenetrating radiation incident
upon a semi-infinite solid with uniform properties, the Thermal sweating. → Sweating, thermal.
value of the thermal inertia for the surface can be deter-
mined in appropriate units by √kpc = 2π–1/2 · t1/2 · E · Thermal tachypnea: A rapid respiratory frequency ac-
∆T–1 in which ∆T is the rise in surface temperature at companied by an increase in respiratory minute volume
time, t. and, commonly, a decrease in tidal volume, in response
to a thermoregulatory need to dissipate heat. (Gk. ta-
Thermal insulation. → Thermal resistance. khus-swift; pnoia-breath) Synonym: thermal polyp-
nea.
Thermal insulation, clothing (Icl): The intrinsic insu-
lation of a clothing assembly. The effective insulation Thermoeffector: An organ system and its action, re-
of clothing is (Icl + Ia) where Ia is the reciprocal of the spectively, that affect heat balance in a controlled
thermal conductance of the ambient environment. (Icl + manner as part of the processes of temperature regu-
Ia) is usually measured as the temperature gradient lation.
from the surface of a heated man-sized manikin to the Note: A multitude of thermoeffectors is involved in
ambient air divided by the heat production per unit area both autonomic and behavioral temperature regula-
of manikin surface. [°C · m2 · W–1] The value is some- tion.
times expressed in clo units.
Thermoeffector gain: The derivative of the thermoef-
Thermal panting: Increased respiratory evaporative fector output with respect to body temperature devia-
heat loss (REHL) due to increased respiratory minute tion from the set-point.
volume. Note:The gain may change due to fever, adaptation
Note: Thermal panting in animals can occur both with and other processes. In combination with thermoeffec-
a closed and open mouth. tor threshold temperature, thermoeffector gain is an
important characteristic of thermoeffectors.
Thermal polypnea. → Thermal tachypnea.
Note: Although polypnea is more commonly used, ta- Thermoeffector threshold: The level of activity of a
chypnea is etymologically more correct. potential thermoeffector that is transgressed when it
becomes actively involved in temperature regulation.
Thermal radiance. → Radiance, thermal. Note: For some of the thermoeffectors, thresholds can
be precisely determined, e.g., as the level of basal met-
Thermal radiant exitance. → Radiant exitance, abolic rate (BMR) or resting metabolic rate (RMR)
thermal. of a tachymetabolic animal above which metabolic
heat production (H) will increase in response to a suf-
Thermal resistance (R): The reciprocal of thermal ficiently strong cold exposure. The thresholds of other
conductance. [°C · m–2 · W–1] or [°C · W–1] Synonym: thermoeffectors are arbitrarily defined, or agreed upon
thermal insulation. by convention, because basal or resting levels are diffi-
cult to define, e.g., in case of circulatory convection of
Thermal shell. → Shell, thermal. heat to the skin and the underlying cutaneous vasomo-
tor tone.
Thermal strain: In temperature regulators: 1. Any
deviation of body temperature induced by sustained Thermoeffector threshold temperature: Describes
thermal stress that cannot be fully compensated by the level of a specified body temperature (e.g., core
temperature or mean body temperature) the trans- nonshivering thermogenesis (NST), observed partic-
gression of which in one direction, either upward or ularly in small mammals, is the brown adipose tissue
downward, will activate a certain thermoeffector. As a (BAT).
rule, the threshold core temperature determined for a
given effector will be a function of skin temperature, Thermogenesis, nonshivering (obligatory)
and vice versa; similar interdependencies exist with and (NST(O)): That component of nonshivering thermo-
among the threshold temperatures determined for spec- genesis (NST) (i.e., heat production unrelated to the
ified thermosensor regions in the body core (e.g., hy- contractions of voluntary muscles) that is independent
pothalamus, spinal cord). Evaluation of mean body of short-term changes in ambient temperature (Ta).
temperature threshold values tries to account for the Note: NST(O) corresponds to basal metabolic rate
entire thermal input from the body. Specified threshold (BMR) or standard metabolic rate (SMR). Although
temperatures for certain thermoeffectors may change NST(O) is unaffected by short-term exposure to cold, it
during the processes of fever, acclimat(izat)ion and may be changed by processes of acclimat(izat)ion to
adaptation. Threshold temperature and thermoeffec- sustained cold or heat stress or feeding conditions.
tor gain are the dominant parameters describing the
thermoeffector output as a function of body tempera- Thermogenesis, nonshivering, (thermoregulatory)
ture, thus, they constitute essential characteristics of the (NST(T)): The increase in nonshivering thermogene-
thermal controller. sis (NST) in response to acute cold exposure. The prin-
cipal effector organ is the brown adipose tissue
Thermoeffector threshold zone (Interthreshold (BAT). which may adaptively increase its capacity for
zone): The temperature range between two threshold heat production in the course of acclimat(izat)ion and
(body) temperatures, for activation of any thermoef- adaptation to cold stress.
fector responses, particularly of metabolic heat pro- Note: The term nonshivering thermogenesis (NST)
duction (H) and of evaporative heat loss when no usually refers to NST(T).
thermal load is present. This special steady-state may
be called set-point. Thermoeffector threshold zone Thermogenesis, shivering: An increase in the rate of
should be distinguished from thermoneutral zone heat production during cold exposure due to increased
(TNZ). contractile activity of skeletal muscles not involving
voluntary movements and external work. [W], [W·
Thermogenesis, diet-induced: Increase of obligatory kg–1], [W · m–2] or [W · m–3]
nonshivering thermogenesis occurring especially in Note: Shivering thermogenesis progresses, as its in-
rodents when transferred from standard food to a highly tensity increases, from thermoregulatory muscle
palatable cafeteria diet, of which the animals consume tone, to micro-vibrations, to clonic contractions of both
more but dissipate part of the surplus caloric intake by flexor and extensor muscles. All shivering thermo-
enhanced heat production. The brown adipose tissue genesis is blocked by curare.
(BAT) is considered as the main effector organ of
diet-induced thermogenesis. Thermogenin. → P 32000 Polypeptide.
Note: The relationship between postprandial (excess)
heat production and diet-induced thermogenesis is not Thermography, infra-red: The recording of the tem-
presently clarified. perature distribution of a body from the infra-red radi-
Thermogenesis, nonshivering (NST): Heat produc- ation emitted by the surface.
tion due to metabolic energy transformation by pro-
cesses that do not involve contractions of skeletal Thermolability, controlled. → Temperature lability,
muscles, i.e., tone, microvibrations, tremor (→ shiver- controlled.
ing), or tonic or voluntary contractions. In thermal Thermolability, passive. → Temperature lability,
physiology this term is conventionally used to indicate passive.
thermoregulatory (cold-induced) nonshivering ther-
mogenesis (NST). [W], [W · m–2], [W · kg–1] Thermoneutral zone (TNZ): The range of ambient
Note: While all organs contribute to obligatory non- temperature at which temperature regulation is
shivering thermogenesis according to their rates of achieved only by control of sensible heat loss, i.e.,
resting metabolism, the principal effector organ re- without regulatory changes in metabolic heat produc-
sponsible for the short-term or adaptive activation of tion (H) or evaporative heat loss. The thermoneutral
zone (TNZ) will therefore be different when insulation, Thermoregulatory nonshivering thermogenesis. →
posture or basal metabolic rate (BMR) vary. The term Nonshivering thermogenesis (thermoregulatory).
thermoneutral zone (TNZ) does not apply to ecto-
therms. The thermoneutral zone (TNZ) should be Thermoresponsive: Neural element which changes its
distinguished from thermoeffector threshold zone. activity in response to changes of its own temperature.
Synonyms: temperature responsive, thermosensi-
Thermopreferendum: The thermal conditions that an tive.
individual organism or a species selects for its ambient Note: In comparison to the more general term temper-
environment in natural or experimental circumstances. ature dependence, neuronal thermoresponsiveness
implies at least the possibility of generation or modula-
Thermoreactive: Descriptive of neural elements tion of information by temperature, either non-specific
whose activity changes with the temperature of a re- or as a specific temperature signal. The synonymous
mote region of the body, due to synaptic input from this term thermosensitive implies the specificity of the
region. thermoresponsive neural element as a temperature sig-
Note: A central nervous neuron may be both thermore- nal generator.
active and thermoresponsive (thermosensitive).
Thermosensitive: Descriptive of thermoresponsive
Thermoreceptor: Thermosensitive neural element neural structures with the implication that the neural el-
for which both its afferent function and its response ements involved provide specific temperature signals.
characteristics are electrophysiologically identified (→ Synonym: temperature sensitive.
thermosensor). Synonym: temperature receptor.
Note: Thermoreceptors have been unequivocally Thermosensor: Neural element or circuitry of neural
identified, so far, only in the skin and mucous surfaces elements for which it is established by psychophysical
as cold receptors, warm receptors, and infrared re- criteria or analysis of thermoeffector responses or
ceptors. changes of core temperature that they transduce tem-
perature in such a way that thermal sensation is elicited
Thermoregulation. → Temperature regulation. and/or temperature regulation is adequately stimu-
lated.
Thermoregulatory behavior. → Temperature regu- Note: Thermosensors may not (yet) be accessible to
lation, behavioral. electrophysiological identification, but their function is
equivalent to that of thermoreceptors which are elec-
Thermoregulatory behavior, natural: → Tempera- trophysiologically identified thermosensors.
ture regulation, behavioral.
Thermotolerance: A rapid, short acting molecular
Thermoregulatory behavior, operant. → Tempera- process associated with the synthesis of several fami-
ture regulation, behavioral. lies of heat shock proteins (HSP) of different molecu-
lar weights elicited as a result of acute short sublethal
Thermoregulatory conditioned reflex: The physio- heat injury. It is thought to protect cells from noxious
logical (autonomic and behavioral) responses of an stimuli as well as to accelerate their repair. It is also de-
organism to changes in its thermal environment, which fined as heat shock response (HSR). The time course
can also be elicited by a conditioned stimulus. of heat shock proteins (HSP) vary in different cells
but, on the average, heat shock proteins (HSP) in the
Thermoregulatory muscle tone: The increase in the intact body seem to operate several hours following the
electrical activity of the skeletal musculature of a rest- stress and retains its activity for a few days. The re-
ing tachymetabolic temperature regulator during sponse is not heat-specific and can be elicited subse-
moderate cooling. During more intensive cooling, quent to subjection to several other stressors (e.g.,
thermoregulatory muscle tone will be superimposed ischemia, some chemicals, etc.). Synonym: heat shock
by microvibrations and eventually shivering tremor. response (HSR).
The corresponding electrical activities can be deter-
mined qualitatively and quantitatively by means of Thermotropism: The turning or movement of a plant
electromyographical recording (→ shivering). or animal in response to a temperature stimulus. (Gk.
therme-heat; trope-turn)
Thigmothermy: The dependence of the core temper- agocytes) when activated by microorganisms or their
ature of an ectothermic animal on the conductive ex- products. The production of tumor necrosis factor
change of heat with its immediate environment, e.g., (TNF-α) is caused by infection, injury and trauma. It
water, air, soil. (Gk. thigma-touch; therme-heat) (as well as other cytokines) produces the acute phase
response. Tumor necrosis factor-α (TNF-α) is con-
Threshold temperature. → Thermoeffector thresh- sidered an important endogenous pyrogen, as are inter-
old temperature. leukin-1 (IL-1), interleukin-6 (IL-6), and some other
cytokines.
Threshold, thermoeffector. → Thermoeffector
threshold. UCP1: Specialized protein with a subunit Mr 32000 of
the mitochondrial inner membrane of brown adipose
Tissue thermal conductance. → Thermal conduc- tissue (BAT); collapses the electro-chemical proton
tance, tissue. gradient generated by respiration and thus, prevents
ATP synthesis; the amount of this mitochondrial com-
Tolerated ambient temperature range: The range of ponent is increased when brown adipose tissue (BAT)
ambient temperature (Ta) within which the body core is in a recruited state. Synonym: thermogenin, uncou-
temperature can be kept, by means of autonomic pling protein, P 32000 polypeptide.
thermoregulatory processes, within certain limits typ-
ical for the species or the individual under consider- UCP2, UCP3: Proteins of similar amino acid sequence
ation. [°C] to UCP1, but found in tissues other than brown adi-
pose tissue (BAT), discussed as being involved in ther-
Torpor: A state of inactivity and reduced responsive- mogenesis
ness to stimuli (e.g., during hibernation, hypother-
mia, or estivation). Ultraviolet. → Radiant energy.
Total body area (Ab). → Area, total body. Uncoupling protein. → P 32000 Polypeptide.
Total environment: All environmental factors that ex- Upper critical temperature. → Critical tempera-
ert an influence on an organism and to which an organ- ture, upper.
ism must be adequately adapted in order to survive (i.e.,
competitors for food sources and predators as well as Upper temperature survival limit. → Temperature
the many components of the physical environment and survival limit, upper.
the climate).
Useful work accomplished. → Work, positive.
Total heat production: The rate of transformation of
chemical energy into heat in an organism (metabolic Vapor pressure (water). → Pressure, water vapor.
heat production (H)) plus any heat flow liberated
within the body resulting from work done on the or- Volume, thermal expansion coefficient of. → Ther-
ganism by an external force (negative work rate). mal expansion coefficient of volume.
[W], [W · m–2], [W · m–3], [W · kg–1]
Note: During positive work and when no work is being Volumetric specific heat. → Specific heat, volumet-
done on or by the organism, total heat production ric.
equals metabolic heat production (H).
Total radiant absorptance. → Radiant absorptance, Wallowing: The thermoregulatory increase in evapora-
total. tive heat loss by spreading an aqueous fluid (e.g., water,
mud, urine) on the body surface.
Transmittance, radiation (τ): The ratio of the radiant
energy transmitted through a body to the total radiation Warm-blooded: The thermal state of an animal that
incident on it. maintains its core temperature considerably higher
than that of the environment when subjected to a low
Tumor necrosis factor-α (TNF-α): Heat labile pro- ambient temperature (Ta). Synonym: tachymeta-
teins produced by immune cells (e.g., mononuclear ph- bolic (preferred). Antonym: cold-blooded.
Note: This maintained temperature gradient between chanical energy, work production, useful work ac-
the organism and its environment is dependent on the complished. Antonym: workrate, negative.
relatively high rate of metabolic heat production (H)
(tachymetabolism) of warm-blooded animals com- Work production. → Work rate, positive.
pared with the low rate of heat production (bradyme-
tabolism) of cold-blooded animals. Thus, the terms Zone of thermal comfort. → Thermal comfort, zone
tachymetabolic and bradymetabolic are preferred to of.
the terms warm-blooded and cold-blooded because
the first pair of terms relates to a more basic physiolog- Zone of thermal indifference. → Thermal indiffer-
ical distinction and because the second pair of terms has ence, zone of.
been used with various meanings not all of which are
consistent with the definitions given here. Zone of thermoneutrality. → Thermoneutral zone.
Warm-blooded is not a synonym of homeothermic,
because the definition of warm-blooded does not spec-
ify the degree of temperature stability consistent with
homeothermy: the core temperatures of some
warm-blooded animals vary considerably either nych-
themerally or seasonally.