Thermoregulation

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Thermoregulation in animals

Ectotherm Endotherm Poikilotherm Homeothermy

Heterothermy Stenotherm Eurytherm Thermolabile
Thermostability Gigantothermy Kleptothermy
Bradymetabolism Tachymetabolism

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Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even
when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts
the surrounding temperature as its own body temperature, thus avoiding the need for internal thermoregulation.
The internal thermoregulation process is one aspect of homeostasis: a state of dynamic stability in an organism's
internal conditions, maintained far from equilibrium with its environment (the study of such processes in
zoology has been called ecophysiology or physiological ecology). If the body is unable to maintain a normal
temperature and it increases significantly above normal, a condition known as hyperthermia occurs. For
humans, this occurs when the body is exposed to constant temperatures of approximately 55 C (131 F), and
any prolonged exposure (longer than a few hours) at this temperature and up to around 75 C (167 F) death is
almost inevitable.[citation needed] Humans may also experience lethal hyperthermia when the wet bulb temperature is
sustained above 35 C (95 F) for six hours.[1] The opposite condition, when body temperature decreases below
normal levels, is known as hypothermia.
Whereas an organism that thermoregulates is one that keeps its core body temperature within certain limits, a
thermoconformer is subject to changes in body temperature according to changes in the temperature outside of
its body at a certain temperature.
It was not until the introduction of thermometers that any exact data on the temperature of animals could be
obtained. It was then found that local differences were present, since heat production and heat loss vary
considerably in different parts of the body, although the circulation of the blood tends to bring about a mean
temperature of the internal parts. Hence it is important to identify the parts of the body that most closely reflect
the temperature of the internal organs. Also, for such results to be comparable, the measurements must be
conducted under comparable conditions. The rectum has traditionally been considered to reflect most accurately
the temperature of internal parts, or in some cases of sex or species, the vagina, uterus or bladder.
Occasionally the temperature of the urine as it leaves the urethra may be of use in measuring body temperature.
More often the temperature is taken in the mouth, axilla, ear or groin.

Some animals undergo one of various forms of dormancy where the thermoregulation process temporarily
allows the body temperature to drop, thereby conserving energy. Examples include hibernating bears and torpor
in bats.

Contents

1 Classification of animals by thermal characteristics

o 1.1 Endothermy versus ectothermy
1.1.1 Ectotherms
1.1.1.1 Ectothermic cooling
1.1.1.2 Ectothermic heating (or minimizing heat loss)
1.1.2 Endothermy
o 1.2 Homeothermy versus poikilothermy
2 In vertebrates
o 2.1 Brain control
o 2.2 In birds and mammals
o 2.3 In humans
2.3.1 In hot conditions
2.3.2 In hot and humid conditions
2.3.3 In cold conditions
2.3.4 Related diseases and syndromes
2.3.5 Human heat output power
3 In plants
4 Behavioral temperature regulation
o 4.1 Hibernation, estivation, and daily torpor
5 Variations in the temperature of human beings and some animals
o 5.1 Normal human temperature
o 5.2 Variations from thermometer placement
o 5.3 Variations associated with development
o 5.4 Variations due to circadian rhythms
o 5.5 Variations due to women's menstrual cycles
o 5.6 Variations due to fever
o 5.7 Variations due to biofeedback
o 5.8 Variations due to other factors
6 Low body temperature increases lifespan
7 Limits compatible with life
o 7.1 Arthropoda
8 See also
9 References
10 Further reading
11 External links

Classification of animals by thermal characteristics

Endothermy versus ectothermy
Thermoregulation in organisms runs along a spectrum from endothermy to ectothermy. Endotherms create most
of their heat via metabolic processes, and are colloquially referred to as warm-blooded. Ectotherms use external
sources of temperature to regulate their body temperatures. They are colloquially referred to as cold-blooded
despite the fact that body temperatures often stay within the same temperature ranges as warm-blooded animals.
Ectotherms

Main article: Ectotherm

Ectothermic cooling

Seeking shade is one method of cooling. Here Sooty Tern chicks are using a Black-footed Albatross chick for
shade.
Vaporization:
o Evaporation of sweat and other bodily fluids.
Convection:
o Increasing blood flow to body surfaces to maximize heat loss.
Conduction:
o Losing heat by being in contact with a colder surface. For instance:
Lying on cool ground.
Staying wet in a river, lake or sea.
Covering in cool mud.
Radiation:
o releasing heat by radiating it away from the body.

Ectothermic heating (or minimizing heat loss)

Convection:
o Climbing to higher ground up trees, ridges, rocks.
o Entering a warm water or air current.
o Building an insulated nest or burrow.
Conduction:
o Lie on hot rock.
Radiation:
o Lie in sun.
o Angle in relation to sun.
o Fold skin to reduce exposure.
o Conceal wing surfaces.
o Expose wing surfaces.
Insulation
o Change shape to alter surface/volume ratio
o Inflate the body

Thermographic image of a snake around an arm

To cope with low temperatures, some fish have developed the ability to remain functional even when the water
temperature is below freezing; some use natural antifreeze or antifreeze proteins to resist ice crystal formation
in their tissues. Amphibians and reptiles cope with heat loss by evaporative cooling and behavioral adaptations.
An example of behavioral adaptation is that of a lizard lying in the sun on a hot rock in order to heat through
conduction.
Endothermy
Main article: Endotherm
An endotherm is an animal that regulates its own body temperature, typically by keeping it at a constant level.
To regulate body temperature, an organism may need to prevent heat gains in arid environments. Evaporation of
water, either across respiratory surfaces or across the skin in those animals possessing sweat glands, helps in
cooling body temperature to within the organism's tolerance range. Animals with a body covered by fur have
limited ability to sweat, relying heavily on panting to increase evaporation of water across the moist surfaces of
the lungs and the tongue and mouth. Mammals like cats, dogs and pigs, rely on panting or other means for
thermal regulation and have sweat glands only in foot pads and snout. The sweat produced on pads of paws and
on palms and soles mostly serves to increase friction and enhance grip. Birds also avoid overheating by gular
fluttering, flapping the wings near the gular (throat) skin, similar to panting in mammals, since their thin skin
has no sweat glands. Down feathers trap warm air acting as excellent insulators just as hair in mammals acts as
a good insulator. Mammalian skin is much thicker than that of birds and often has a continuous layer of
insulating fat beneath the dermis. In marine mammals, such as whales, or animals that live in very cold regions,
such as the polar bears, this is called blubber. Dense coats found in desert endotherms also aid in preventing
heat gain such as in the case of the camels.
A cold weather strategy is to temporarily decrease metabolic rate, decreasing the temperature difference
between the animal and the air and thereby minimizing heat loss. Furthermore, having a lower metabolic rate is
less energetically expensive. Many animals survive cold frosty nights through torpor, a short-term temporary
drop in body temperature. Organisms when presented with the problem of regulating body temperature have not
only behavioural, physiological, and structural adaptations but also a feedback system to trigger these
adaptations to regulate temperature accordingly. The main features of this system are stimulus, receptor,
modulator, effector and then the feedback of the newly adjusted temperature to the stimulus. This cyclical
process aids in homeostasis.

Homeothermy versus poikilothermy

Homeothermy and poikilothermy refer to how stable an organism's temperature is. Most endothermic
organisms are homeothermic, like mammals. However, animals with facultative endothermy are often
poikilothermic, meaning that their temperature can vary considerably. Most fish are ectotherms, as all of their
heat comes from the surrounding water. However, most are homeotherms because their temperature is very
stable.

In vertebrates
By numerous observations upon humans and other animals, John Hunter showed that the essential difference
between the so-called warm-blooded and cold-blooded animals lies in observed constancy of the temperature of
the former, and the observed variability of the temperature of the latter. Almost all birds and mammals have a
high temperature almost constant and independent of that of the surrounding air (homeothermy). Almost all
other animals display a variation of body temperature, dependent on their surroundings (poikilothermy).
Certain mammals are exceptions to this rule, being warm-blooded during the summer or daytime, but coldblooded during the winter when they hibernate or at night during sleep. J. O. Wakelin Barratt has demonstrated
that, under certain pathological conditions, a warm-blooded (homeothermic) animal may become temporarily
cold-blooded (poikilothermic). He has shown conclusively that this condition exists in rabbits suffering from
rabies during the last period of their life, the rectal temperature being, then, within a few degrees of the room

temperature and varying with it. He explains this condition by the assumption that the nervous mechanism of
heat regulation has become paralysed. The respiration and heart-rate being also retarded during this period, the
resemblance to the condition of hibernation is considerable. Again, Sutherland Simpson has shown that during
deep anaesthesia a warm-blooded animal tends to take the same temperature as that of its environment. He
demonstrated that, when a monkey is kept deeply anaesthetized with ether and is placed in a cold chamber, its
temperature gradually falls, and that, when it has reached a sufficiently low point (about 25 C in the monkey),
the employment of an anaesthetic is no longer necessary, the animal then being insensible to pain and incapable
of being roused by any form of stimulus; it is, in fact, narcotised by cold, and is in a state of what may be called
"artificial hibernation." Once again this is explained by the fact that the heat-regulating mechanism has been
interfered with. Similar results have been obtained from experiments on cats.

Brain control
Thermoregulation in both ectotherms and endotherms is controlled mainly by the preoptic area of the anterior
hypothalamus.[2] Such homeostatic control is separate from the sensation of temperature.[2]

In birds and mammals

Kangaroo licking its arms to cool down on a very hot day

In cold environments, birds and mammals employ the following adaptations and strategies to minimize heat
loss:
1. Using small smooth muscles (arrector pili in mammals), which are attached to feather or hair shafts; this
distorts the surface of the skin making feather/hair shaft stand erect (called goose bumps or pimples)
which slows the movement of air across the skin and minimizes heat loss.
2. Increasing body size to more easily maintain core body temperature (warm-blooded animals in cold
climates tend to be larger than similar species in warmer climates (see Bergmann's Rule))
3. Having the ability to store energy as fat for metabolism
4. Have shortened extremities
5. Have countercurrent blood flow in extremities - this is where the warm arterial blood travelling to the
limb passes the cooler venous blood from the limb and heat is exchanged warming the venous blood and
cooling the arterial (e.g., Arctic Wolf[3] or penguins[4][5])

In warm environments, birds and mammals employ the following adaptations and strategies to maximize heat
loss:
1.
2.
3.
4.

Behavioural adaptations like living in burrows during the day and being nocturnal
Evaporative cooling by perspiration and panting
Storing fat reserves in one place (e.g., camel's hump) to avoid its insulating effect
Elongated, often vascularized extremities to conduct body heat to the air

In humans

Simplified information processing structure of human thermoregulation.

As in other mammals, thermoregulation is an important aspect of human homeostasis. Most body heat is
generated in the deep organs, especially the liver, brain, and heart, and in contraction of skeletal muscles.[6]
Humans have been able to adapt to a great diversity of climates, including hot humid and hot arid. High
temperatures pose serious stresses for the human body, placing it in great danger of injury or even death. For
humans, adaptation to varying climatic conditions includes both physiological mechanisms resulting from
evolution and behavioural mechanisms resulting from conscious cultural adaptations.[7][8]
There are four avenues of heat loss: convection, conduction, radiation, and evaporation. If skin temperature is
greater than that of the surroundings, the body can lose heat by radiation and conduction. But, if the
temperature of the surroundings is greater than that of the skin, the body actually gains heat by radiation and
conduction. In such conditions, the only means by which the body can rid itself of heat is by evaporation. So,
when the surrounding temperature is higher than the skin temperature, anything that prevents adequate
evaporation will cause the internal body temperature to rise.[9] During sports activities, evaporation becomes the
main avenue of heat loss.[10] Humidity affects thermoregulation by limiting sweat evaporation and thus heat
loss.[11]
The skin assists in homeostasis (keeping different aspects of the body constant, e.g. temperature). It does this by
reacting differently to hot and cold conditions so that the inner body temperature remains more or less constant.
Vasodilation and sweating are the primary modes by which humans attempt to lose excess body heat. The brain
creates much heat through the countless reactions which occur. Even the process of thought creates heat. The
head has a complex system of blood vessels, which keeps the brain from overheating by bringing blood to the
thin skin on the head, allowing heat to escape. The effectiveness of these methods is influenced by the character
of the climate and the degree to which the individual is acclimatized.
In hot conditions

Play media
A dog panting after exercise
1. Eccrine sweat glands under the skin secrete sweat (a fluid containing mostly water with some dissolved
ions), which travels up the sweat duct, through the sweat pore and onto the surface of the skin. This
causes heat loss via evaporative cooling; however, a lot of essential water is lost.
2. The hairs on the skin lie flat, preventing heat from being trapped by the layer of still air between the
hairs. This is caused by tiny muscles under the surface of the skin called arrector pili muscles relaxing
so that their attached hair follicles are not erect. These flat hairs increase the flow of air next to the skin
increasing heat loss by convection. When environmental temperature is above core body temperature,
sweating is the only physiological way for humans to lose heat.
3. Arteriolar vasodilation occurs. The smooth muscle walls of the arterioles relax allowing increased blood
flow through the artery. This redirects blood into the superficial capillaries in the skin increasing heat
loss by convection and conduction.
Note: Most animals cannot sweat efficiently. Cats and dogs have sweat glands only on the pads of their feet.
Horses and humans are two of the few animals capable of sweating. Many animals pant rather than sweat
because the lungs have a large surface area and are highly vascularised. Air is inhaled, cooling the surface of the
lungs and is then exhaled losing heat and some water vapour.
In hot and humid conditions
In general, humans appear physiologically well adapted to hot dry conditions.[12] However, effective
thermoregulation is reduced in hot, humid environments such as the Red Sea and Persian Gulf (where
moderately hot summer temperatures are accompanied by unusually high vapor pressures), tropical
environments, and deep mines where the atmosphere can be water-saturated.[12][7] In hot-humid conditions,
clothing can impede efficient evaporation.[8] In such environments, it helps to wear light clothing such as cotton,
that is pervious to sweat but impervious to radiant heat from the sun. This minimizes the gaining of radiant
heat, while allowing as much evaporation to occur as the environment will allow. Clothing such as plastic
fabrics that are impermeable to sweat and thus do not facilitate heat loss through evaporation can actually
contribute to heat stress.[11]
In cold conditions
1. Sweat production is decreased.
2. The minute muscles under the surface of the skin called erector pili muscles (attached to an individual
hair follicle) contract (piloerection), lifting the hair follicle upright. This makes the hairs stand on end,
which acts as an insulating layer, trapping heat. This is what also causes goose bumps since humans do
not have very much hair and the contracted muscles can easily be seen.
3. Arterioles carrying blood to superficial capillaries under the surface of the skin can shrink (constrict),
thereby rerouting blood away from the skin and towards the warmer core of the body. This prevents
blood from losing heat to the surroundings and also prevents the core temperature dropping further. This
process is called vasoconstriction. It is impossible to prevent all heat loss from the blood, only to reduce
it. In extremely cold conditions, excessive vasoconstriction leads to numbness and pale skin. Frostbite
occurs only when water within the cells begins to freeze. This destroys the cell causing damage.
4. Muscles can also receive messages from the thermo-regulatory center of the brain (the hypothalamus) to
cause shivering. This increases heat production as respiration is an exothermic reaction in muscle cells.
Shivering is more effective than exercise at producing heat because the animal remains still. This means
that less heat is lost to the environment through convection. There are two types of shivering: lowintensity and high-intensity. During low-intensity shivering, animals shiver constantly at a low level for
months during cold conditions. During high-intensity shivering, animals shiver violently for a relatively

short time. Both processes consume energy, however high-intensity shivering uses glucose as a fuel
source and low-intensity tends to use fats. This is a primary reason why animals store up food in the
winter.[citation needed]
5. Mitochondria can convert fat directly into heat energy, increasing the temperature of all cells in the
body. Brown fat is specialized for this purpose, and is abundant in newborns and animals that hibernate.
The process explained above, in which the skin regulates body temperature is a part of thermoregulation. This is
one aspect of homeostasis the process by which the body regulates itself to keep internal conditions constant.
Related diseases and syndromes

Hypothermia
Hyperthermia
Heat stroke
Raynaud's phenomenon (Raynaud's disease)
Endocrine System Disorders (Hyperthyroidism, Hypothyroidism)
Induced hypothermia
Erythromelalgia (hyperthermia)

Human heat output power

An average-size human will output from around 70 watts to 870 watts, depending on the amount of physical
activity undertaken.[13]

In plants
Thermogenesis occurs in the flowers of many plants in the Araceae family as well as in cycad cones.[14] In
addition, the Sacred lotus (Nelumbo nucifera) is able to thermoregulate itself,[15] remaining on average 20 C
(36 F) above air temperature while flowering. Heat is produced by breaking down the starch that was stored in
their roots,[16] which requires the consumption of oxygen at a rate approaching that of a flying hummingbird.[17]
One possible explanation for plant thermoregulation is to provide protection against cold temperature. For
example, the skunk cabbage is not frost-resistant, yet it begins to grow and flower when there is still snow on
the ground.[14] Another theory is that thermogenicity helps attract pollinators, which is borne out by observations
that heat production is accompanied by the arrival of beetles or flies.[18]

Behavioral temperature regulation

Animals other than humans regulate and maintain their body temperature with physiological adjustments and
behavior. Desert lizards are ectotherms and so unable to metabolically control their temperature but can do this
by altering their location. They may do this by in the morning only raising their head from its burrow and then
exposing their entire body. By basking in the sun, the lizard absorbs solar heat. It may also absorb heat by
conduction from heated rocks that have stored radiant solar energy. To lower their temperature, lizards exhibit
varied behaviors. Sand seas, or ergs, produce up to 136 F (57.7C), and the sand lizard will hold its feet up in the
air to cool down, seek cooler objects with which to contact, find shade or return to their burrow. They also go to
their burrows to avoid cooling when the sun goes down or the temperature falls.

During cold weather western bluebirds can increase their thermal inertia by huddling.
Animals also engage in kleptothermy in which they share or even steal each other's body warmth. In
endotherms such as bats[19] and birds (such as the mousebird[20] and emperor penguin[21]) it allows the sharing of
body heat (particularly amongst juveniles). This allows the individuals to increase their thermal inertia (as with
gigantothermy) and so reduce heat loss.[22] Some ectotherms share burrows of ectotherms. Other animals exploit
termite mounds.[23][24]
Some animals living in cold environments maintain their body temperature by preventing heat loss. Their fur
grows more densely to increase the amount of insulation. Some animals are regionally heterothermic and are
able to allow their less insulated extremities to cool to temperatures much lower than their core temperature
nearly to 0 C. This minimizes heat loss through less insulated body parts, like the legs, feet (or hooves), and
nose.

An ostrich can keep its body temperature very constant, even though it can be very hot during the day and cold
at night.

Hibernation, estivation, and daily torpor

To cope with limited food resources and low temperatures, some mammals hibernate during cold periods. To
remain in "stasis" for long periods, these animals build up brown fat reserves and slow all body functions. True
hibernators (e.g., groundhogs) keep their body temperatures low throughout hibernation whereas the core
temperature of false hibernators (e.g., bears) varies; occasionally the animal may emerge from its den for brief
periods. Some bats are true hibernators and rely upon a rapid, non-shivering thermogenesis of their brown fat
deposit to bring them out of hibernation.
Estivation is similar to hibernation but usually occurs in hot periods to allow animals to avoid high
temperatures and desiccation. Both terrestrial and aquatic invertebrate and vertebrates enter into estivation.
Examples include Lady beetles (Coccinellidae),[25] North American desert tortoises, crocodiles, salamanders,
cane toads,[26] and the Water-holding Frog[27]
Daily torpor occurs in small endotherms like bats and hummingbirds, which temporarily reduces their high
metabolic rates to conserve energy.[28]

Variations in the temperature of human beings and some animals

Chart showing diurnal variation in body temperature, ranging from about 37.5 C from 10 a.m. to 6 p.m., and
falling to about 36.3 C from 2 a.m. to 6 a.m.

Normal human temperature

Main article: Normal human body temperature
Previously, average oral temperature for healthy adults had been considered 37.0 C (98.6 F), while normal
ranges are 36.1 C (97.0 F) to 37.8 C (100.0 F). In Poland and Russia, the temperature had been measured
axillary. 36.6 C was considered "ideal" temperature in these countries, while normal ranges are 36 C to 36.9
C.
Recent studies suggest that the average temperature for healthy adults is 98.2 F or 36.8 C (same result in three
different studies). Variations (one standard deviation) from three other studies are:

36.4 - 37.1 C (97.5 - 98.8 F)

36.3 - 37.1 C (97.3 - 98.8 F) for males, 36.5 - 37.3 C (97.7 - 99.1 F) for females
36.6 - 37.3 C (97.9 - 99.1 F)[29]

Variations from thermometer placement

Temperature varies according to thermometer placement, with rectal temperature being 0.3-0.6 C (0.5-1 F)
higher than oral temperature, while axillary temperature is 0.3-0.6 C (0.5-1 F) lower than oral temperature.[30]
The average difference between oral and axillary temperatures of Indian children aged 612 was found to be
only 0.1 C (standard deviation 0.2 C),[31] and the mean difference in Maltese children aged 414 between oral
and axillary temperature was 0.56 C, while the mean difference between rectal and axillary temperature for
children under 4 years old was 0.38 C.[32]

Variations associated with development

Of the lower warm-blooded animals, there are some that appear to be cold-blooded at birth. Kittens, rabbits,
and puppies, if removed from their surroundings shortly after birth, lose their body heat until their temperature
has fallen to within a few degrees of that of the surrounding air. But such animals are at birth blind, helpless,
and in some cases furless. Animals that are born when in a condition of greater development can maintain a
fairly constant body temperature. In strong, healthy human infants a day or two old, the temperature rises
slightly when removed, but in that of weakly, ill-developed children it either remains stationary or falls. The
cause of the variable temperature in infants and young immature animals is the imperfect development of the
nervous regulating mechanism.
The average temperature falls slightly from infancy to puberty and again from puberty to middle age, but after
that stage is passed the temperature begins to rise again, and by about the eightieth year is as high as in infancy.

Variations due to circadian rhythms

In humans, a diurnal variation has been observed dependent on the periods of rest and activity, lowest at 11 p.m.
to 3 a.m. and peaking at 10 a.m. to 6 p.m. Monkeys also have a well-marked and regular diurnal variation of
body temperature that follows periods of rest and activity, and is not dependent on the incidence of day and

night; nocturnal monkeys reach their highest body temperature at night and lowest during the day. Sutherland
Simpson and J.J. Galbraith observed that all nocturnal animals and birds - whose periods of rest and activity are
naturally reversed through habit and not from outside interference - experience their highest temperature during
the natural period of activity (night) and lowest during the period of rest (day). Those diurnal temperatures can
be reversed by reversing their daily routine.[33]
In essence, the temperature curve of diurnal birds is similar to that of man and other homoeothermal animals,
except that the maximum occurs earlier in the afternoon and the minimum earlier in the morning. Also, the
curves obtained from rabbits, guinea pigs, and dogs were quite similar to those from man. These observations
indicate that body temperature is partially regulated by circadian rhythms.

Variations due to women's menstrual cycles

During the follicular phase (which lasts from the first day of menstruation until the day of ovulation), the
average basal body temperature in women ranges from 36.45 to 36.7 C (97.6 to 98.1 F). Within 24 hours of
ovulation, women experience an elevation of 0.15 - 0.45 C (0.2 - 0.9 F) due to the increased metabolic rate
caused by sharply elevated levels of progesterone. The basal body temperature ranges between 36.7 - 37.3C
(98.1 - 99.2F) throughout the luteal phase, and drops down to pre-ovulatory levels within a few days of
menstruation.[34] Women can chart this phenomenon to determine whether and when they are ovulating, so as to
aid conception or contraception.

Variations due to fever

Fever is a regulated elevation of the set point of core temperature in the hypothalamus, caused by circulating
pyrogens produced by the immune system. To the subject, a rise in core temperature due to fever may result in
feeling cold in an environment where people without fever do not.

Variations due to biofeedback

Some monks are known to practice Tummo, biofeedback meditation techniques, that allow them to raise their
body temperatures substantially.[35]

Variations due to other factors

In Simpson's & Galbraith's work, the mean temperature of the female was higher than that of the male in all the
species examined whose sex had been determined.
Meals sometimes cause a slight elevation, sometimes a slight depressionalcohol seems always to produce a
fall. Exercise and variations of external temperature within ordinary limits cause very slight change, as there are
many compensating influences at work, which are discussed later. The core temperature of those living in the
tropics is within a similar range to those dwelling in the Arctic regions.

Low body temperature increases lifespan

It was long theorised that low body temperature may prolong life. On November 2006, a team of scientists from
the Scripps Research Institute reported that transgenic mice that had body temperature 0.3-0.5 C lower than
normal mice (due to overexpressing the uncoupling protein 2 in hypocretin neurons (Hcrt-UCP2), which
elevated hypothalamic temperature, thus forcing the hypothalamus to lower body temperature) indeed lived
longer than normal mice. The lifespan was 12% longer for males and 20% longer for females. Mice were
allowed to eat as much as they wanted.[36][37][38] The effects of such a genetic change in body temperature on
longevity is harder to study in humans. The UCP2 genetic alleles seen in humans so far are associated with
obesity[39]

Limits compatible with life

There are limits both of heat and cold that an endothermic animal can bear and other far wider limits that an
ectothermic animal may endure and yet live. The effect of too extreme a cold is to decrease metabolism, and
hence to lessen the production of heat. Both catabolic and anabolic pathways share in this metabolic depression,
and, though less energy is used up, still less energy is generated. The effects of this diminished metabolism
become telling on the central nervous system first, especially the brain and those parts concerning
consciousness;[40] both heart rate and respiration rate decrease; judgment becomes impaired as drowsiness
supervenes, becoming steadily deeper until the individual loses consciousness; without medical intervention,
death by hypothermia quickly follows. Occasionally, however, convulsions may set in towards the end, and
death is caused by asphyxia.[40]
In experiments on cats performed by Sutherland Simpson and Percy T. Herring, the animals were unable to
survive when rectal temperature fell below 16C.[40] At this low temperature, respiration became increasingly
feeble; heart-impulse usually continued after respiration had ceased, the beats becoming very irregular,
appearing to cease, then beginning again. Death appeared to be mainly due to asphyxia, and the only certain
sign that it had taken place was the loss of knee-jerks.
However, too high a temperature speeds up the metabolism of different tissues to such a rate that their
metabolic capital is soon exhausted. Blood that is too warm produces dyspnea by exhausting the metabolic
capital of the respiratory centre;[citation needed] heart rate is increased; the beats then become arrhythmic and
eventually cease. The central nervous system is also profoundly affected by hyperthermia and delirium, and
convulsions may set in. Consciousness may also be lost, propelling the person into a comatose condition. These
changes can sometimes also be observed in patients suffering from an acute fever.[citation needed] The lower limit of
temperature that humans can endure depends on many factors, but no one can survive a temperature of 45 C
(113 F) or above for very long.[citation needed] Mammalian muscle becomes rigid with heat rigor at about 50C,
with the sudden rigidity of the whole body rendering life impossible.
H.M. Vernon has done work on the death temperature and paralysis temperature (temperature of heat rigor) of
various animals. He found that species of the same class showed very similar temperature values, those from
the Amphibia examined being 38.5C, Fish 39C, Reptilia 45C, and various Molluscs 46C.[citation needed] Also, in
the case of pelagic animals, he showed a relation between death temperature and the quantity of solid
constituents of the body. In higher animals, however, his experiments tend to show that there is greater variation
in both the chemical and physical characteristics of the protoplasm and, hence, greater variation in the extreme
temperature compatible with life.

Arthropoda
The maximum temperatures tolerated by certain thermophilic arthropods exceeds the lethal temperatures for
most mammals, birds, reptiles, fish and amphibians.[41]
The most heat-resistant insects are three genera of desert ants recorded from three different parts of the world.
The ants have developed a lifestyle of scavenging for short durations during the hottest hours of the day, in
excess of 50 C (122 F) and often approaching 70 C (158 F), for the carcasses of insects and other forms of
life which have succumbed to heat stress.[42]
In April 2014, the South Californian mite Paratarsotomus macropalpis has been recorded as the world's fastest
land animal relative to body length, at a speed of 322 body lengths per second. Besides the unusually great
speed of the mites, the researchers were surprised to find the mites running at such speeds on concrete at
temperatures up to 60 C (140 F), which is significant because this temperature is well above the lethal limit
for the majority of animal species. In addition, the mites are able to stop and change direction very quickly.[41]