Temperature Regulation Dr.

Sudipta Chahraborty
Assistant Professor of Zoology
Maulana Azad College, Kolkata

Metabolic Rate:
The basal metabolic rate (BMR) is the stable rate of energy metabolism measured in mammals and
birds under conditions of minimum environmental and physiological stress (namely, at rest with no
temperature stress) and after fasting has temporarily halted digestive and absorptive processes.

 Ambient temperature affects body temperature in almost all animals other than birds and mammals.
 Since the minimum metabolic rate varies with the body temperature, it is necessary to measure the
equivalent of the basal metabolic tare at a controlled, specified body temperature at which the animal is
not expending additional metabolic energy to warm or cool itself.
 The standard metabolic rate (SMR) is defined as an animal's resting and fasting metabolism at a given
body temperature.
 Interestingly, the SMR of some ectotherms depends on their previous temperature history, owing to
metabolic compensation or thermal acclimation, which is described later.

Remark:
 BMR and SMR give little information about the metabolic costs of normal activities carried out by the
animals because for these measurements, the animal is in an unnaturally controlled and quiet state.
 The best term to describe the metabolic rate of an animal in its natural state is called its field metabolic
rate (FMR), which is the average rate of energy utilization as the animal goes about its normal
activities, which may range from complete inactivity during resting to maximum exertion when
chasing prey (or being preyed upon).

Metabolic Scope
The range of metabolic rates of which an animal is capable is called its aerobic metabolic scope, defined as the
ratio of the maximum sustainable metabolic rate to the BMR (or the SMR) determined under controlled,
resting conditions.
 This metabolic scope values like 5, 7, or 14 indicates the increase in an animal's maximal energy
expenditure (usually measured as oxygen consumption) over and above the amount that it expends
under resting conditions.
 Commonly, metabolic rate increases from 10 to 15 times with activity in many animals.
 Since the sustained activity is normally powered by aerobic metabolism, this type of measurement does
not take into account the contribution of anaerobic processes to activity.
 The processes build up an oxygen debt and hence are not sustainable.

Figure: An oxygen deficit develops during a period of sustained, intense activity. Active muscle tissue with
anaerobic capabilities can amass oxygen deficit, which is subsequently paid off in the form of a delayed oxidation of
an anaerobic product such as lactic acid. As a result, an elevated metabolic rate continues after cessation of activity
but gradually subsides with time. The initial oxygen deficit is due to the use of preexisting stores of high-energy
phosphagens amassed during rest. Replenishment of these stores is included in the repayment of the oxygen debt.
Specific Dynamic Action
Max Rubner reported in 1885 that a marked increase in metabolism accompanies the processes of
digestion and assimilation of food independently of other activities-- a phenomenon named specific dynamic
action (SDA).
 Since then, SDA has been documented in all five vertebrate classes, as well as in invertebrates including
crustaceans, insects, and molluscs.
 Generally, an animal's oxygen consumption and heat production increases within about 1 hour after a
meal is eaten, reaching a peak some 3-6 hours later and remaining elevated above the basal value for
several hours.
 In fish, amphibians, and reptiles with an SDA equivalent to a doubling or tripling of metabolic rate,
there are also attendant large increases in heart rate and cardiac output and a temporary redistribution
of blood toward the gut.
 Similar cardiovascular changes of lesser magnitude occur in animals with less prominent SDA
responses (e.g., in humans).
 The mechanism of SDA is presumably related to the fact that certain organs, such as the liver, expend
extra energy processing recently absorbed nutrients for entry into metabolic pathways. The extra
energy consumed by such processes is lost as heat. The increase in heat production differs, depending
on the ingested food materials. The magnitude of the increased metabolic rate ranges from 5% to 10%
of total energy of ingested carbohydrates and fats and from 25% to 30% of that of proteins.

Figure: Specific dynamic artion occurs

after feeding In the toad Bufo marinus.
Specific dynamic action was induced by
injecting peptone (a mixture of amino
acids produced from chemically digested
meat protein) into the animal's stomach.

Body Size and Metabolic Rate

The study of how both anatomical and physiological characteristics change with body mass is called scaling. Changes in
body size introduce changes that are not always simple and in geometric proportion.
 For example, the doubling of the height of an animal while retaining the same body proportions is accompanied
by a fourfold increase in surface area and an eightfold increase in mass.
Changes in body mass have great effects on an animal's metabolic rate.
 For example, the respiratory and metabolic requirements of a tiny water shrew during diving compared with
those of a submerged whale. Although both whales and water shrews normally dive, a whale can hold its breath
and remain under water far longer than a shrew.
The reason stems from the general principle that:
“small animals must respire at higher rates per unit body mass than large animals”.
In fact, there is an inverse relation between the rate of oxygen consumption per gram of body mass and the total mass of
the animal.
 For example, a 100 gram mammal consumes far more energy per unit mass per unit time than does a 1000 gram
mammal.
Mass-specific metabolic rate:
It is a reality that, area is proportional to the square of a linear dimension while volume is proportional to the
cube of the linear dimension. Further, doubling of the height of an animal while retaining the same body
proportions results in fourfold increase in surface area and an eightfold increase in mass. The consequences of
non-geometric scaling for the functional anatomy and physiology of the animal are immediately evident.
Metabolic rate is a power function of body mass, as described by the simple relation:
MR = aMb, in which,
MR is the basal or standard metabolic rate, M is the body mass, a is the intercept of the log-log regression line
(and differs between species), and b is an empirically determined exponent that expresses the rate of change of
MR with change in body mass.

Mass-specific metabolic rate, also termed metabolic intensity, is the metabolic rate of a unit mass of tissue (i.e.,
amount of oxygen, consumed per kilogram per hour). It is determined by dividing both sides of the abpve
equation by M:
MR = aMb = aM(b-1)
M M ,
As such, when overall metabolic rate rises with increasing body mass, whereas the mass-specific metabolic
rate (metabolic rate of a unit mass of tissue) decreases with increasing body mass. This principle first emerges
in the mouse-to-elephant plot as depicted below:

(a) (b)
Figure: Generalised relationship between mass and (a) metabolic rate (b) mass-specific metabolic rate. Mass-
specific metabolic rate (MR) in mammals declines as body mass (M) increases (A) The mouse-to-elephant
curve, with rnetabolic intensity (mass-specific metabolic rate) given as O2 consumption per unit mass plotted
against body mass.

To attain linear plot in logarithmic form, we get,

log MR = log a + b (log M)
or, log MR log a + (b-1) log M, which when plotted stands as:
,

Figure: log-log plot og mass-specific metabolic rate equation

Respiratory Quotient
To translate the amount of oxygen consumed into equivalent heat production, we must know the
relative amounts of carbon and hydrogen oxidized. The oxidation of hydrogen atoms is hard to determine,
however, because metabolic water (i.e., that produced by oxidation of hydrogen atoms available in foodstuffs),
together with other water, is lost in the urine and from a variety of body surfaces at a rate that is irregular and
determined by unrelated factors (e.g., osmotic stress and ambient relative humidity). The ratio of the volume
of CO2, produced to the volume of O2, removed from within a given time is called respiratory quotient (RQ):

Under resting, steady-state conditions, the RQ in Table below, is characteristic of the type of molecule
catabolized (carbohydrate, fat, or protein). Thus, the RQ reflects the proportions of carbon and hydrogen in the
food molecules. The following examples illustrate how the RQ of the major food types may be calculated from
a formulation of their oxidation reactions:

For Carbohydrates:
The general formula of carbohydrates is (C6H12O6). In the complete oxidation of a carbohydrate, oxygen
is used in effect only to oxidize the carbon to form CO2. Upon complete oxidation, each mole of a carbohydrate
produces n moles of both H2O and CO2, and consumes n moles of O2. The RQ for carbohydrate oxidation is
thus 1. The overall catabolism of glucose, for example, may be formulated as:

,
For Fats:
As different fats contain different ratios of carbon, hydrogen, and oxygen, they differ slightly in their
RQs. The RQ characteristic of the oxidation of a fat such as tripalmitin may be calculated as follows:

For Protein:
 The RQ characteristic of protein catabolism is a bit complex because proteins are not completely broken
down in oxidative metabolism.
 Some of the oxygen and carbon of the constituent amino acid residues remains combined with nitrogen
and is excreted as nitrogenous wastes in urine and feces.
 In mammals, the excreted end product is urea, (NH2)2CO; in birds, it is primarily uric acid, C5H4N4O2.
 To obtain the RQ, it is therefore necessary to know the amount of ingested protein as well as the amount
and kind of nitrogenous wastes excreted.
 The oxidation of carbon and hydrogen in the catabolism of protein typically produces

Remark:
It is routinely assumed in making deductions from RQ that:
 the only substances metabolized are carbohydrates, fats, and proteins;
 no synthesis takes place alongside breakdown; and
 the amount of CO2, exhaled in a given time equals the CO2, produced by the tissues in that interval.
(a) (b)
Figure: Open and closed respirometery can be combined in a single experiment to measure an animal's gas exchange
partitioning between various sites. In this experiment on the reed-fish Calamoicthys calabaricus, two independent open
systems are used to determine branchial and cutaneous aquatic oxygen uptake. A third, closed respirometery system
includes the air in the funnel above the head, from which the animal breathes air. Gas samples taken after an air breath
are used to calculate aerial oxygen consumption. Results depicted in figure (b)

Max Rubner’s theory of surface hypothesis

Max Rubner proposed that the metabolic rate of birds and mammals that maintain a more or less constant
body temperature should be proportional to body surface area because the rate of heat transfer between two
compartments (i.e., warm animal body and cool environment) is proportional, all else being equal, to their area
of mutual contact.

Isometry:
 The surface area of an object of isometric shape (i.e., nonvarying proportions) and uniform density
varies as the 0.67 (or f) power of its mass.
 This is because mass increases as the cube of linear dimension, whereas surface area increases only as
the square.
 As noted, this relation holds true for a series of animals of different mass only if body proportions
remain constant.
 This provision is generally satisfied only by adult individuals of different size within a species, because
they tend to obey the principle of isometry -namely, proportionality of shape regardless of size. In this
case, it follows that the surface area must vary as the 0.67 power of body mass.

Figure: Relationship amongst dimensions of geometrically similar objects

 Figure: Geometric similarity in inanimate and animal world

Allometry:
 However, the principle of isometry is not followed in individuals of different size belonging to related
but different species.
 Instead, they tend to follow the principle of allometry- namely, systematic changes in body proportions
with increasing species size.
 An example of allometry was alluded to earlier when we compared the proportions of an elephant
with those of a mouse.
 In a comparison of surface- to-mass relations in mammals of different species ranging from mice to
whales, surface areas were found to be proportional to the 0.63 power of body mass

Figure: Geometric dissimilarity in inanimate and animal world

Remark:
In spite of the logical attractiveness of the surface hypothesis, it is not without flaws.
 True, the difference in metabolic intensity between large and small homeotherms may indeed be an
adaptation to the more rapid loss of heat from a smaller animal owing to surface-to-volume relations,
the small animal having more surface area per unit mass.
 Another flaw of the surface hypothesis arises from the simple observation that the metabolic rates of
animals whose body temperatures vary with that of their surroundings (such as fish, amphibians,
reptiles and most invertebrates) exhibit nearly the same relation to body mass as the metabolic rates of
animals that actively maintain a constant, high body temperature (i.e., birds and mammals.
 Scaling effects are also evident at the cellular level. There is a correlation between differences in
metabolic intensity of animals of differing sizes and the number of mitochondria per unit volume of
tissue. The cells of a small mammal contain more mitochondria and mitochondria1 enzymes in a given
volume of tissue than do the cells of a large mammal. Because mitochondria are sites of oxidative
respiration, this correlation comes as no surprise
Homeoviscous adaptation
Acclimatization of ectothermic animals to cold or hot environments in major is due to the fact that the
membrane lipids become more saturated during acclimatization to warmth and less saturated during
acclimatization to cold, helping to stabilize the form of the lipids and thus the cellular functions that spring
from them. This phenomenon is called homeoviscous adaptation, referring to adaptations at the molecular
level through natural selection that help minimize temperature-induced differences in viscosity.

Experimental proof:
 Most often used as an index is the steady-state fluorescence anisotropy (a measure of the lack of symmetry of a
molecule or structure)
 1,6-Diphenyl-l,3,5-hexatriene (DPH) ) is a commonly used probe of membrane fluidity.
 A high fluorescence anisotropy indicates a high degree of lipid polarization and membrane order attendant with
a low membrane viscosity.
 Result shows changes in DPH polarization of the basolateral membranes of enterocytes isolated from rainbow
trout.
 Initially, acute temperature changes are accompanied by changes in membrane polarization and fluidity.
 However, with time, homeoviscous adaptation of the membrane lipds results in a lipid polarization and
membrane viscosity after acclimation at 5°C that are similar to those after acclimation at 20°C.

Figure: Homeoviscous adaptation maintains relatively constant membrane lipid properties in rainbow trout enterocytes.
After an initial measurement at 25'C (point I), a warm-acclimated (20°C) trout is rapidly cooled to 5°C. Initially, the
membranes of its enterocytes become more polarized and more viscous (point 2), but as homeoviscous adaptation oc-
curs the lipid membranes become less polarized and regain their fluid- ity (point 3). Similarly, if a cold-acclimated trout at
a measurement tem- perature of 5°C is rapidly warmed to 25°C and measured, the lipid membranes are initially highly
depolarized (point 4) but become more polarized with acclimation (point 1). [Adapted from Hazel, 1995.]

Remark:
Exceptionally, some animals become fully acclimated to temperature change with moderate or even no homeoviscous
adaptation in lipid membrane properties. Altered expression of membrane proteins and proliferation of mitochondria1
and sarcoplasmic reticular membranes, along with homeoviscous adaptation of membrane lipids, present a picture of cell
membranes as dynamic structures that change in complex ways to retain their function de- spite temperature change.

Finally, there are also regional differences in lipid prop- erties, including melting point, in some mammals. In the limbs,
which may be subjected to near-freezing tempera- tures, the tissue lipids are less saturated and so have a lower melting
point than the fats in the body core.
Determinants of Body Heat and Temperature
The temperature of an animal depends on the amount of heat (calories) contained per unit mass of tissue. Since
tissues consist primarily of water, the heat capacity of tissues between 0°C and 40°C approximates 1.0 cal. 0C -1.g-1. It
follows that the larger the animal, the greater its body heat content at a given temperature. The rate of change of body
heat depends on
(1) the rate of heat production through metabolic means,
(2) the rate of external heat gain, and
(3) the rate of heat loss to the environment
We can state that,
body heat = heat produced + (heat gained - heat lost)
= heat produced + heat transferred
Thus, body heat, and hence the body temperature of an animal, can be regulated by changes in the rate of heat production
and heat transfer or exchange (i.e., heat gained minus heat lost).
The total heat content of an animal is determined by the metabolic production of heat and the thermal flux
between the animal and its terrestrial surroundings. The relation between these factors can be represented as:
Htot = Hv + Hc + Hr + He + Hs, where,
Htot = total heat
Hv = heat produced metabolically
Hc = heat lost or gained by convection and conduction
Hr = net heat transfer by radiation
He = heat lost by evaporation
Hs = heat stored in the body

Figure: Processes of heat is transferred between an animal and the environment.

Conduction:
The transfer of heat between objects and substances that are in contact with each other is conduction. It
results from the direct transfer of kinetic energy of the motion from molecule to molecule, with the net flow of energy
being from the warmer to the cooler region. The rate of heat transfer through a solid conductor of uniform properties can
be expressed as:

,
in which Q is the rate of heat transfer (in joules per centimeter per second) by conduction; k is the thermal
conductivity of the conductor; A is the cross-sectional area (in square centimeters); and 1 is the distance (in
centimeters) between points 1 and 2, which are at temperatures t1 and t2,respectively.
 Conduction is not limited to heat flow within a given substance; it may also be between two phases,
such as the flow of heat from skin into the air or water in contact with the body surface.

Radiation
The transfer of heat by electromagnetic radiation takes place without direct contact between objects. All
physical bodies at a temperature above absolute zero emit electromagnetic radiation in proportion to the
fourth power of the absolute temperature of the surface. As an example of how radiation works, the sun's rays
may warm a black body to a temperature well above the temperature of the air surrounding the body. A dark
body both radiates and absorbs more strongly than does a more reflective body having a lower emissivity. For
temperature differences between the surfaces of two bodies of about 20 Celsius degrees or less, the net radiant
heat exchange is approximately proportional to the temperature difference.

Evaporation
Every liquid has its own latent heat of vaporization, which is the amount of energy required to change it
from a liquid to a gas of the same temperature-that is, to evaporate. The energy required to convert 1 g of
water into water vapor is relatively high, approximately 585 cal. Many animals dissipate heat by allowing
water to be evaporated from body surfaces.

Heat storage:
Heat storage leads to an increase in temperature of the heat-storing mass. The larger the mass, or the
higher its specific heat, the smaller its rise in temperature (in 0C) for a given quantity of heat (in joules)
absorbed. Thus, a large animal that has a small surface-to-mass ratio tends to heat up more slowly in response
to an environmental heat load than does a small animal that has a relatively high surface-to-mass ratio.

The rate of heat transfer (kilocalories per hour) into or out of an animal also depends on several factors.
Changing the value of any one of them alters heat flow across the body surface in the direction of the
temperature gradient:

 Surface area per gram of tissue decreases with increases in body mass, providing small animals with a
high heat flux per unit of body weight (as already noted). Animals can sometimes control their
apparent surface area by changing posture (e.g., by extending limbs or drawing them close to body).
 Temperature difference between the environment and the animal's body has a large effect by altering
the temperature gradient (change in temperature per unit distance) for heat transfer. The closer an animal
maintains its temperature to the ambient temperature, the less heat will flow into or out of its body.
 Specific heat conductance of the animal's surface varies with the nature of the body surface. Animals
with high heat conductances in surface tissues are typically close to the temperature of their
surroundings, with some exceptions, such as the elevation of body temperature when an animal basks
in sunlight. Animals that actively maintain a constant body temperature (birds, mammals) have
feathers, fur, or blubber that decrease the heat conductance of their body surfaces

Figure: Blood flow to the skin helps regulate the heat conductance of the body surface. Vasomotor control of
peripheral arterioles shunts the arterial blood either to the skin or away from it.
  In response to environmental cold, peripheral blood vessels vasoconstrict, shunting blood away from
the surface of an endotherm.
 In response to high temperatures, the blood is diverted to the skin, where it approaches temperature
equilibrium with the environment.
In ectotherms, cutaneous blood flow is often increased through peripheral vasodilation to absorb heat from the
environment.)
Animals use several different mechanisms to regulate the exchange of heat between themselves and the
environment:
 Behavioral control includes moving to a part of the environment where heat exchange with the
environment favors attaining optimal body temperature. For instance, a desert ground squirrel retires
to its burrow during the midday heat; a lizard suns itself to gain heat by radiation from its
surroundings, raising its body temperature well above ambient temperature. Animals also control the
amount of surface area available for heat exchange by adjusting their postures.
 Autonomic control of blood flow to the vertebrate skin affects the temperature gradient and, hence, the
heat flux at the body surface. For example, the activation of piloerector muscles increases the extent of
fluffing of pelage and plumage, which increases the effectiveness of insulation by increasing the
amount of trapped, unstirred air. Sweating and salivation during panting cause evaporative cooling.
 Acclimatization includes long-term changes in pelage or subdermal, fatty-layer insulation, as well as
changes in the capacity for autonomic control of evaporative heat loss through sweating.

Figure: Fur and blubber act as heat insulation. (A) Fur is outside the skin and circulation, and its insulating properties
can be changed rapidly only by flattening or fluffing through pilomotor control. (B) Because blubber is located under the
skin and is supplied with blood vessels, its insulating value can be regulated by shunting the blood through vasomotor
control to the surface or away from the surface below the blubber

Poikilotherms
They are those animals in which body temperature tends to fluctuate more or less with the ambient
temperature when air or water temperatures are varied experimentally.

Homeotherms
They (or homoiotherms) maintain body temperatures above ambient temperatures and regulate their
body temperatures within a narrow physiological range by controlling heat production and heat loss. In most
mammals, the normal physiological range for core body temperature is typically from 37°C to 38°C; whereas,
in birds, it is closer to 40°C.

Ectotherms
They produce metabolic heat at comparatively low rates-rates normally too low to allow for
endothermy. Often, ectotherms have low rates of metabolic heat production and high thermal conductances-
that is, they are poorly insulated. As a result, heat derived from metabolic processes is quickly lost to cooler
surroundings.
Endotherms
They are animals that generate their own body heat through heat production as a by-product of
metabolism, typically elevating their body temperatures considerably above ambient temperatures.
 Most produce heat metabolically at high rates, and many have relatively low thermal conductivity
because of good insulation (fur, feathers, fat), which enables them to conserve heat in spite of a high
temperature gradient between body and environment.
 Mammals and birds exemplify animals that regulate their temperatures within relatively narrow limits
and are therefore said to be homeothermic endotherms.
 A few large fishes (sharks and larger tuna) and some flying insects are termed regional heterothermic
endotherms because they maintain regions of their body above ambient temperatures, sometimes for
short periods of time under specific circumstances, as in flying insects.

Heterotherms are those animals capable of varying degrees of endothermic heat production, but they
generally do not regulate body temperature within a narrow range. They may be divided into two groups,
regional and temporal heterotherms.
 Temporal heterotherms constitute a broad category of animals whose temperatures vary widely over
time. Monotremes (egg-laying mammals) such as the echidna are temporal heterotherms, as are other
mammals and birds in torpor and hibernation.
 Regional heterotherms are generally ectotherms that can achieve high core (i.e., deep-tissue)
temperatures through muscular activity, while their peripheral tissues and extremities approach the
ambient temperature. Examples include mako sharks, tuna, and many flying insects. Elevated
temperatures generally allow higher metabolic rates than would be achieved at ambient environmental
temperatures. Fishes that are regional heterotherms depend on countercurrent heat exchangers.

(a) (b) (c)

Figure: (a) Homeotherms maintain body temperature as ambient temperature changes, whereas the body temperatures
of poikilotherms more closely track ambient temperature (b) The relation between body temperature and ambient
temperature differs in various animals. The cat is a strict homeotherm, maintaining body temperature independent of
ambient temperature, whereas monotremes (platypus and echidna) are temporal heterotherms. The lizard is a strict
heterotherm. (c) Endotherms can be regionally heterothermlc in mammals

Metabolic control of locusts while preparing for flight

Some flying insects, including locusts, beetles, cicadas, and arctic flies, can be considered both temporal and
regional heterotherms because, when preparing to fly, they raise the core temperatures in their thoracic parts
to more or less regulated levels.
Event:
 At moderate ambient temperatures, these insects are unable to take off and fly without prior warmup
because their flight muscles contract too slowly to produce sufficient power for flight at temperatures
much below 40°C.
 When inactive, however, these insects behave strictly as ectotherms.
 After such an insect is aloft, its flight muscles produce enough heat to maintain elevated muscle
temperatures, and the in- sect even employs heat-dissipating mechanisms to prevent overheating.
Explanation
 These flying insects generally have quite a large mass; and some, such as bumblebees, butterflies, and moths, are
covered with heat-insulating "hairs" or scales.
 To warm up, these insects activate their large thoracic flight muscles, which are among the most metabolically
active tis- sues known.
 The activation is such that antagonistic muscles work against one another, producing heat without much wing
movement other than small, rapid vibrations akin to shivering.
 Flight is finally initiated when the thoracic temperature has reached the temperature that is main tained during
flight, about 40°C

Figure: The sphinx moth Manduca sexta undergoes a pre-flight thermogenesls. Shiverlng of the thoracic flight muscles
causes a steep Increase In thoracic temperature prior to flight [Adapted from Heinrich, 1974.1]

Ground squirrel regulate metabolic output during hibernation

A period of deep torpor, or winter dormancy, hibernation lasts for weeks or even several months in cold climates. It is
entered into through slow wave sleep and is devoid of rapid-eye-movement sleep. Hibernation is common in mammals
of the orders Rodentia, Insectivora, and Chiroptera, which can store sufficient energy reserves to sur- vive the periods of
nonfeeding.
 During hibernation, the hypothalamic thermostat is reset to as low as 20 Celsius degrees or more below normal.
 At ambient temperatures between 5°C and 150C, many hibernators keep their temperatures as little as 1 degree
above ambient temperature.
 If the air temperature falls to dangerously low levels, the animal increases its metabolic rate to maintain a
constant low Tb or becomes aroused.
For ground squirrel which was kept in a chamber having a temperature of 4°C
:
 Metabolism increases briefly during an episode of arousal from hibernation in a ground squirrel.
 At onset of hibernation, the set point for temperature is depressed.
 Metabolism decreases, allowing T, to drop to 1-3 Celsius degrees above T, throughout hibernation.
 Arousal occurs when the set-point temperature climbs to 38'C, and a strong surge of metabolic heat production
raises Tb to the new set-point level. Abbreviation: RAMR, resting average metabolic rate. [From Swan, 1974]

Figure: Experiment judging hibernation in ground squirrel

Supercooling:
Some animals can undergo supercooling, in which the body fluids can be cooled to below their freezing
temperature, yet remain unfrozen because ice crystals fail to form. Ice crystals will not form if they have no nuclei to
initiate crystal formation.
 Certain fishes dwelling at the bottom of Arctic fjords live in a continually supercooled state and normally do not
freeze. If they brush against frozen ice on the water surface, however, ice crystallizes rapidly throughout their
bodies and they die immediately.
 Thus, survival for these fish depends on their remaining well below the surface where ice is absent. The body
fluids of some cold-climate ectotherms contain antifreeze substances.
 For example, the body fluids of a number of arthropods, including mites and various insects, contain glycerol, the
concentration of which typically increases in the winter. Glycerol, acting as an antifreeze solute, lowers the
freezing point to as low as - 17°C. The tissues of larvae of the parasitic wasp Brachon cephi can withstand even
lower temperatures; they have been supercooled to -47°C without ice crystal formation.
The blood of the antarctic ice fish Trematomus contains a glycoprotein antifreeze that is from 200 to 500 times as effective
as an equivalent concentration of sodium chloride in preventing ice crystal formation. The glycoprotein lowers the
temperature at which the ice crystals enlarge, but it does not lower the temperature at which they melt.

Thermogenesis
When the ambient temperature drops below the lower critical temperature, an endothermic animal responds by
generating large amounts of additional heat from energy stores, thereby preventing a decrease in the core temperature.
There are two primary means of extra heat production other than exercise: shivering and nonshivering thermogenesis.
Both processes convert chemical energy into heat by a normal energy-converting metabolic mechanism that is adapted to
primarily produce heat. Essentially all the chemical-bond energy released in this process is fully degraded to heat rather
than to chemical or mechanical work. Shivering is a means of using muscle contraction to liberate heat.

 Shivering thermogenesis occurs in some insects as well as in endothermic vertebrates. The nervous system
activates groups of antagonistic skeletal muscles so that there is little net muscle movement other than shivering.
 The activation of muscle causes ATP to be hydrolyzed to provide energy for contraction.
 As the muscle contractions are inefficiently timed and mutually opposed, they produce no useful physical work,
but the chemical energy released during contraction appears as heat.
 In nonshivering thermogenesis, enzyme systems for the metabolism of fats are activated throughout the body, so
conventional fats are broken down and oxidized to produce heat.
 Very little of the energy released is conserved in the form of newly synthesized ATP.
 A specialization found in a few mammals for fat-fueled thermogenesis is brown fat, also called brown adipose
tissue (BAT).
 Generally found as small deposits in the neck and between the shoulders brown fat is an adaptation for rapid,
massive heat production.
 This fat contains such extensive vascularization and so many mitochondria that it is brown (owing largely to
mitochondrial cytochrome oxidase) rather than white.
 In ordinary body fat, deposits must first be reduced to fatty acids, which enter the circulation and eventually are
taken up by other tissues, where they are oxidized. In brown fat, oxidation takes place within the fat cells
themselves, which are richly endowed with fat-metabolizing enzyme systems.

The mechanism:
 Nonshvering thermogenesis in fat (including brown fat) is activated by the sympathetic nervous system through
the release of norepinephrine, which binds to receptors on the adipose cells of brown-fat tissue.
 Through a second-messenger mechanism, this signal leads to thermogenesis by two mechanisms. In the first of
these mechanisms, normal ATP utilization for cellular processes rises in these fat cells in response to the
sympathetic signal, accounting for part of the increased heat production.
 Through processes such as ion pumping by the plasma membrane, ATP is hydrolyzed to produce work and heat.
 In the second mechanism, ATP production is uncoupled during respiratory chain oxidation. The resynthesis of
ATP from ADP and Pi is normally coupled to the movement of protons (H +) down their electrochemical gradient
from intermembrane space into mitochondria across the inner mitochondrial membrane.
 Thermogenesis in brown fat is characterized by the appearance in the inner mitochondrial membrane of
uncoupling proteins, which provide a pathway for protons to leak across this membrane without the energy of
their downhill movement being harnessed for the phosphorylation of ADP to ATP.
  Once inside the mitochondrion, the protons oxidize substrate oxygen to produce water and heat, or else further
utilization of metabolic energy is required to subsequently pump them into the intermembrane space and
eventually out of the mitochondria.

Hypothalamus as a thermostat
Standing outside in cold temperature condition drops your skin temperature quickly. This stimulates skin cold
receptors (increase in their activity) and cools the blood flowing into the skin. These signals are received by both the
hypothalamic thermostat and higher cortical centers. The thermostat is also activated by the change in blood temperature.
It initiates responses that promote heat gain and inhibits centers that promote heat loss. The activation of Sympathetic
Centers results in several responses including
1) Norepinephrine release from sympathetic fibers constricts skin vessels.
2) Brown fat (found in infants and some animals) oxidation increases causing thermogenesis.
3) Piloerection, occurs which traps air close to skin.
4) Epinephrine secretion from adrenal medulla increases thermogenesis.
A Shivering Center in the hypothalamus is also activated which activates the Brainstem Motor Centers to initiate
involuntary contraction of skeletal muscles causing shivering, which generates heat. Cold also activates some
compensatory behavioral responses including huddling, voluntary physical activity (hand rubbing, pacing), sheltering
next to a heat source and wearing warm clothing. Voluntary or semivoluntary behaviors in response to cold are activated
by the higher brain centers, mainly the cortex and limbic system. When the environmental temperature decreases
gradually (ex. summer to fall), the hypothalamus releases Thyrotropin Releasing Hormone which activates the anterior
pituitary gland to release Thyroid Stimulating Hormone (TSH). TSH induces the thyroid gland to liberate large amounts
of thyroid hormone (T3 and T4) into the blood. Thyroid hormone increases metabolic rate, which increases the amount of
body heat production. As the body gets warmer, the hypothalamic sensors detect the warmth and diminish the heat
producing and heat loss prevention responses.