ANIMAL PHYSIOLOGY HANDOUT

PHYSIOLOGY DEFINED: establishment of space-time relationships between physical and chemical events
within the organism as a response to events both within the organism and in the environment.

MAJOR SUB-DIVISIONS of animal physiology

1. CELL PHYSIOLOGY: it deals with the events that occur within cells and the function of a cell per se. It
also is concerned with the coordinated activities of cells. It tends to be strongly based in biochemistry.

2. ORGAN or WHOLE ORGANISM PHYSIOLOGY: which deals primarily with the way large groups of
different cells (organs) interact with other organs or with how the entire organism behaves a
physiological unit.

Specific Approaches to Animal Physiology:

1. Comparative Physiology: studies the underlying physiological similarities and differences of

organisms.

2. Physiological Ecology: primarily concerned with understanding and describing the physiological
mechanisms that allow an organism to live in a particular environment.

3. Mammalian (human) Physiology: emphasis is on medically-related problems, this is the best funded
and most workers are in this area. As a result, many of the basic discoveries in physiology occur in this
field.

CONTROL SYSTEMS

In order to understand regulation, we need to understand control systems. A control system

consists of a series of components that work together to control some process. It is important to realize
that several components may in some cases reside in a single cell while in other cases many different
cells or even tissues may make up one part of a control system.

1. CONTROLLED VARIABLE: the factor that we wish to regulate. Some type of SENSOR monitors the
value of the controlled variable and sends this information to an error detector.

2. SET POINT: the desired value of the controlled variable. As an example, for alert but not active
humans the set point for body temperature is 37º C. The value of the set point is often relayed to a place
called the error detector as a REFERENCE SIGNAL.

3. ERROR DETECTOR: a device that compares the value of the controlled variable with the set point. It
produces a signal (ERROR SIGNAL) that then operates one or more EFFECTOR SYSTEMS.

ENERGETICS: the study of how energy transformations take place and how these affect the organism.

THERMODYNAMICS:
1. First law: matter/energy is neither created nor destroyed in any process, it simply changes form. (The
law of conservation of matter/energy).

Thus, we know that:

a. For mass: the total amount of material that enters an organism must equal that which leaves or which
remains a part of the organism.
b. Likewise, for energy: the total energy obtained by food or via some physical means (e.g. sunlight,
heat) must either remain stored in the organism or be lost to the environment.

2. Second Law: in any SPONTANEOUS PROCESS the entropy of the UNIVERSE must increase. By entropy,
of course, we mean disorder.

METABOLISM is usually viewed as the sum of two processes: CATABOLISM and ANABOLISM (+
movement).

a. Catabolism -- reactions that break down complex compounds often with the purpose of energy
release with the goal of temporarily conserving some of the released energy in compounds such as ATP.
b. Anabolism and Movement: reactions that use energy to join relatively simple compounds into more
complex ones (ex: protein synthesis, DNA synthesis, etc) or which produce motion.

 Catabolic processes are thermodynamically favorable, anabolic and movement processes are
not.

METABOLISM AND ITS MEASUREMENT:

Metabolism is indicative of the magnitude of at least three very important physiological factors:

1. GROWTH AND REPRODUCTIVE RATE (i.e., rates of synthesis)

2. ACTIVITY -- energy needed to operate the various biochemical pathways needed directly and
indirectly in mechanical locomotion.
3. COMPLEXITY, since presumably the more complex the organism, the greater the metabolism required
to maintain that complexity.

HOW DO WE MEASURE METABOLISM?

Energy: Metabolic Cost: Nearly all chemical reactions involve changes in stored energy (potential
energy, PE) such as the release or absorption of heat or some other type of energy. The total heat
released by an organism can be used as an estimate of an organism's metabolism; measure metabolic
cost as heat production, with units either of calories or Joules.

Power: Metabolic Rate: the metabolic rate has units of power -- usually watts or calories per time.

METHODS USED TO DETERMINE METABOLIC RATE or METABOLISM:

a. The "WHAT COMES IN MUST GO OUT" method - sum all of the PE contained in materials that go in
and out of an organism:

b. HEAT PRODUCTION -- Direct Calorimetry

 Encase the organism in a calorimeter and measure the heat production. The heat production is
proportional to the total metabolic rate since the doing of chemical work results in heat production as a
by-product.

THE MEASUREMENT OF METABOLISM AND METABOLIC RATE USING OXYGEN CONSUMPTION AND
CARBON DIOXIDE PRODUCTION

Metabolism can be measured in aerobic organisms by recording O2 consumption and CO2

production. This requires that the organism must, as a whole be acting aerobically when the
measurements are taken. Thus, if any anaerobic metabolism is occurring in some tissues, its effects must
be reversed elsewhere in the organism.

For most animals, this will mean that if any anaerobic by product is being produced (for example,
lactic acid) it must be used up (not eliminated) by some other part of the body. If anaerobic products
build up (increase) or if they are eliminated from the body, then the entire energy demand cannot be
estimated just from anaerobic processes.

The ATP cycle. Anabolic reactions and movement (muscle action, active transport, exocytosis, etc.) are
non-spontaneous processes. Thus, they all require some sort of energy source and that source is usually
ATP. These processes that need energy from ATP is referred to as "DEMAND" reactions.

The reactions that take energy from complex, energy storage molecules such as carbohydrates or fats
and transfer some of that energy to the formation of ATP (or more simply ~P) is referred to as the
"SUPPLY" reactions.

The rate of the demand reactions is set by some factor (for instance the need to grow or move) and
the supply must meet it. ATP is used to shuttle energy from the energy-rich compounds degraded in the
supply reactions to those needing energy in the demand side. The supply reactions must be adjusted to
keep the level of ATP roughly constant.

Aerobic Metabolism and the RQ Concept*

In most animals, most of the supply reactions responsible for the generation of ~P occur in the
mitochondria (e.g., Krebs cycle, oxidative phosphorylation, β− oxidation of fatty acids). They are closely
related to other reactions that occur in the cytosol (glycolysis and some types of protein and lipid
metabolism).

1. In aerobic organisms most ~P comes from the mitochondria.

KREBS CYCLE (TCA): degrades 2C (acetyl) compounds into lots of high-energy electrons and some ~P.

Nearly all of the energy is removed from the fuel molecules in the form or high-energy electrons. The
carriers that actually move the electrons are the coenzymes NAD+ or NADoxdized and FAD. When these
are reduced by the addition of electrons, they become NADH (NADreduced) and FADH2 respectively.
The energy in these electrons is ultimately extracted by the electron transport system (ETS) and is used
to produce ~P by a process called oxidative phosphorylation.
 Some ~P is also produced by a process called substrate-level phosphorylation. Substrate-level
phosphorylation can be defined as the process where a ~ P is created on a substrate molecule and is
then transferred to some other molecule -- for instance ADP or GDP to make ATP or GDP. In the case of
the Krebs cycle, the substrate-level phosphorylation involves the production of GTP. A single ~P is
produced by substrate level phosphorylation per 2C that enters the Krebs Cycle.

The rate of the Krebs Cycle is controlled by:

1. availability of appropriate 2C compounds

2. availability of electron acceptors (NAD+ and FAD).
3. the concentrations of certain enzyme modulators such as ADP and ATP.

The Krebs cycle gets its fuels from one of three sources:

1. Carbohydrate, 2C fragments after processing in glycolysis (cytosol) and the mitochondrial "bridge
reaction (PDH complex rx)".
2. Fatty acids via β-oxidation in the mitochondria -- this feeds 2 C fragments in to the cycle at the same
place as carbs enter.
3. Sometimes (rarely in humans) by oxidation of the skeletons of amino acids -- these are basically what
are left of an amino acid after it has been deaminated (after the -NH2 is removed).

* CO2 is the Krebs cycle waste product.

The CYTOCHROME ELECTRON TRANSPORT SYSTEM (ETS) :

(i) The ETS accepts high-energy electrons produced by the Krebs cycle and by other mitochondrial and
cytosol reactions such as glycolysis and β-oxidation.
(ii) These electrons are eventually transferred in an orderly manner from one carrier to the next until
they eventually combine with oxygen and form water.
(iii) In the process, energy from the electrons is used to form ATP from ADP and Pi by a process involving
pumping and re-entry of H+.
(iv) The amount of energy that can be conserved in ~P depends on the source of the electrons. Electrons
from NADH are more energetic than those from FADH2 .

The ETS is able to conserve 3 ~P per electron pair from NADH (one pair NADH) and 2 ~P per FADH2
(one pair of electrons per FADH2).

The ETS is controlled by:

1. Availability of O2.
2. Availability of NADH or FADH2.
3. Ratio of ATP to ADP –ADP must be present to allow the system to work.

Keeping track of conserved energy: Per 2C that entered the Krebs cycle, we got:

(a) 1 ~P directly by substrate level phosphorylation

(b) 3 NADH, each of which will give up their electrons to the ETS and produce 3 ~P -- so the total amount
of ~P made from the NADH yielded in one turn of the Krebs cycle is 3* 3 = 9.
(c) there was 1 FADH2 which will give a pair of electrons to the ETS and yield 2 ~P.
(d) Total is 12 ~P.

GLYCOLYSIS: a system that is resident in the cytosol. It takes glucose (and other substances) and
converts to them into 3C fragments. Energy-wise, ATP is generated from ADP and Pi via substrate-level
phosphorylations and high-energy electrons are produced and captured by NAD+. However, these can
only be used to synthesize ~P under aerobic conditions.

The input is always a hexose sugar or hexose derivative. It can either be glucose (or a number of
other hexoses) or glycogen. If we start from glucose, two ~P are expended in preparing the molecule for
the degradative parts of glycolysis. If we start from glycogen, only one is needed because a hexose is
cleaved from glycogen using a Pi.

(a) per hexose, we get two molecules of "waste" at the bottom. Under aerobic conditions, this waste is
pyruvate (the conjugate base of pyruvic acid). Under aerobic conditions, the pyruvate will go the
mitochondria and be completely oxidized in the "bridge reaction" and the Krebs cycle.

(b) per hexose, there are a total of 4 substrate level phosphorylations (i.e., two per pyruvate).

(c) per hexose, there are two molecules of NADH produced from NAD+. Under aerobic conditions, the
electrons on these NADH will eventually end up in the ETS.

So, here is the summary:

Energy Gains:

1. From substrate-level phosphorylations -- a total of 4.

2. 2 NADH (two pairs of high energy electrons) – the number varies between 2 and 3 depending on the
cell type. In muscles it is usually 2 and in the liver it can be 3.

* We only get 2 ~P from oxidative phosphorylation for each NADH produced in glycolysis. That gives a
total of 4 ~P from oxidative phosphorylation.

3. Gross ~P yield = 4 (substrate level) + 4 (ox-phos) = 8 ~P total .

Energy costs: if our hexose was glucose, we paid 2 ~P and if it was glycogen our immediate cost was 1~P
(we paid the other one earlier in synthesizing the glycogen). Thus, costs are 1 or 2 ~P.

Net Yield: Gross - "cost": for glucose, 8 - 2 = 6 ~P , from glycogen, 8 -1 = 7 ~P.

Wastes: two 3-carbon molecules (pyruvate).

Getting from Glycolysis to the Krebs Cycle -- the so-called "bridge reaction" (PYRUVATE
DEHYDROGENASE COMPLEX REACTION OR THE PDH RX FOR SHORT)

If the Krebs cycle is going, and there is adequate NAD+ and FAD, it will be able to accept more 2 C
fragments. Pyruvate, the "waste" product of aerobic glycolysis has 3 C. To use it in the Krebs cycle, we
must remove one carbon. This is done by the "bridge reaction" -- more properly, the pyruvate
dehydrogenase reaction; this occurs in the mitochondrion.

Energetics of the "bridge reaction": the NADH molecules produced by the pyruvate kinase reaction
will yield 3 ~P each since they are produced within the mitochondria. Since the pyruvate dehydrogenase
(a.k.a. bridge) reaction will occur twice per hexose that enters glycolysis (since we get two pyruvates),
then a total of 2 NADH are yielded which gives us a total of 6 ~P.

How much ~P is yielded per hexose?

1. From glycolysis: 6 or 7
2. From the "bridge": 6
3. From the Krebs cycle and ETS: 24
4. TOTAL: 36 or 37 per hexose
5. waste products -- 6 CO2 and 6 H2O

ANAEROBIC METABOLISM

ANAEROBIC METABOLISM: where the ultimate electron acceptor is something different than oxygen.

Glycolysis can be either aerobic or anaerobic, in the sense that if there is adequate O2, electrons
produced in glycolysis enter the mitochondria and the ETA and eventually are attached to O2 and H+ to
form water. Thus, in aerobic metabolism, O2 is the ultimate electron acceptor.

RELATIVE OR ABSOLUTE LACK OF O2:

Relative: More electrons are being produced by the Krebs cycle and glycolysis than can be accepted by
the ETS. There may be large amounts of O2 present, they are simply not large enough to accept all the
electrons being produced. If no alternative acceptor can be found for these electrons, the cell will run
out of the oxidized form of NAD and this will decrease or stop glycolysis and the Krebs cycle. ~P
production will be reduced or stopped.

Absolute: Here, there is no O2 present (anoxia) and therefore the ETS cannot accept electrons. Since
the Krebs cycle is highly dependent on the ETS, it will stop, but it is still possible for glycolysis to proceed
if there are alternative means of storing these electrons until later.

The crucial step in making glycolysis anaerobic is to find an alternative acceptor for the NADH
produced in glycolysis. There are many ways this is done. Here are the two most famous:

(a) the production of lactic acid (lactate): Lactate is simply reduced pyruvate. One reason for the
common use of this particular pathway is that pyruvate production is the terminal step in glycolysis,
therefore using it as an alternative electron acceptor in anaerobic glycolysis allows the entire process
to be self-contained.

(b) the production of ethanol: In the normal activities of an animal, there are many situations where
anaerobic metabolism is significant. The most common situations are heavy exercise or any prolonged
exercise in animals that are primarily "designed" for burst type activities.
 The role of anaerobic glycolysis in the former group is to provide higher ~P production (metabolism)
rates than can be sustained aerobically. This need typically occurs under high workload conditions such
as sprinting.

Examples of latter would be many reptiles, amphibians, fish and spiders -- animals that primarily
capture prey by ambush or quick motion. These animals are also called "sit and wait" predators. If they
are forced to exercise for long periods of time, they rely on anaerobic metabolism as they simply lack
well developed aerobic pathways.

What happens to the lactic acid?

1. The side effects of high [lactic acid]: the principal side effect of lactic acid is that it lowers cellular pH.
This affects the conformation of all proteins within the cell. Generally the cells can tolerate a degree of
pH shift, but large amounts of lactic acid accumulate the pH change will be large enough to significantly
affect the structures of a broad range of proteins. The result is the reduced function that we call fatigue.
There are also psychological effects of high [lactic acid] -- we generally experience such concentrations
as a burning sensation in active muscles.

2. Removal of lactic acid: lactic acid is not a waste in animals. That is because it contains abundant
energy (remember it is reduced pyruvic acid and pyruvic acid is normally oxidized in the Krebs cycle for
energy). However, the cells that make lactic acid typically need to get rid of it because when they make
it they are doing so because, they lack sufficient O2 to oxidize it and also because of the pH problems
just mentioned. And so, like in bacteria, the cells simply dump the lactate -- in this case into the blood
where it quickly is diluted and therefore causes less of a problem. As to what happens next, that
depends:

(a) The heart can oxidize lactate to CO2 and water. Lactate is a preferred fuel of the heart muscle. The
heart usually has no trouble dealing with lactic acid since it has abundant O2. It handles it by reversing
the LDH reaction to get NADH and pyruvate. The NADH is oxidized by the ETS to give ATP and the
pyruvate proceeds through the Krebs cycle as usual. In this case, glycolysis starts in some muscle tissue
and produces lactic acid which is then oxidized in the heart. The net result is aerobic glycolysis where
the first steps occur in the muscle (the anaerobic steps) and aerobic completion occurs in the heart. The
heart may get 50% of more of its energy from lactic acid during exercise.

(b) GLUCONEOGENESIS. In this case, the lactic acid is picked up by the liver (or sometimes other
tissues). It is converted back to glucose at a net cost of energy. Gluconeogenesis is an aerobic process. It
generally proceeds at a slow rate. When associated with exercise, gluconeogenesis occurs mostly during
the recovery period after the exercise. Gluconeogenesis is an umbrella term for a number of processes
that produce glucose from other compounds. Starting points can also be a number of amino acids, Krebs
cycle intermediates, or fatty acids.

Since the accumulation of anaerobic products is associated with fatigue, activity with an appreciable
anaerobic component can only be sustained for a limited duration of time. Activities with appreciable
anaerobic components are referred to as NONSTEADY STATE ACTIVITIES. Here it refers to whether or
not the exercise can continue.
The Regulation of Cellular Metabolism*

Some Important Factors Which Determine Reaction Rates in Biological Systems

Flux is the rate at which material passes through a pathway or any of its steps (the term is also used to
discuss the turnover of energy or movement of substances).

B. Enzyme activity:

"A"ase
A -----------------> B

Enzyme activity is a term that refers to the amount of functional enzyme present (here the enzyme
"A"ase) in some tissue or solution. It is measured in terms of the amount of product produced per unit
time (flux) under ideal conditions and is usually normalized to the amount of protein present (not all the
protein will be the enzyme of interest) or to the amount of tissue the enzyme was removed from.

The most important single factor is the amount of functional enzyme present (also known as the
enzyme activity). Most reactions in organisms are increased by a factor on the order of 107 over the
same reaction outside of the body. When compared under the same conditions of temperature,
pressure, pH, and the same chemical environment, the magnitude of the increase depends simply on
the availability of functional catalyst -- the more enzyme, the faster the reaction rate.

The functional amount that is present is determined by gene expression and by various factors that
modulate the functionality of the enzyme molecules that are present. These include both specific
modulators and chemical and physical factors such as pH, ionic concentration, and temperature.
Temperature, independent of its effect on enzyme structure has its usual role in accelerating reactions.

Enzyme Catalyzed Reactions in Pathways.

1. Equilibrial reactions are the most common type -- here the functional enzyme is in sufficient quantity
that it converts reactant into product very fast. The result is that the reaction is always close to
equilibrium regardless of the flux. Flux through any equilibrial reaction is largely determined by the
availability of reactants and the removal of products, not by regulating the functionality of the
enzyme.

2. Non-equilibrial reactions are rarer but are extremely important in regulating the flux through
pathways. They are called non-equilibrial because at least some of the time the mass action ratio is
very far from equilibrium. The enzyme catalyzing this step is regulated into a shape that makes it a poor
catalyst. As a result, reactants accumulate due to the action of reactions upstream from the non-
equilibrial step and meanwhile reactions that are downstream drain off much of the product.

TEMPERATURE REGULATION: ECTOTHERMY*

What are the effects of temperature on an animal (or plant)?

 There are several answers to this question, some have to do with direct effects on rates of reaction,
others with avoidance of physical damage to membranes and proteins, others with points that make
optimal use of energy available for certain processes.

Three terms relating to the temperature range an organism tolerates should be learned:

1. Stenothermic: tolerates only a narrow range of temperatures: ex. many tropical and deep-sea
organisms.
2. Eurythermal: tolerates a wide range of temperatures.

 The two terms above may be applied to both daily swings in temperature and also to seasonal
changes.

3. Eccritic temperature: the preferred temperature.

LETHAL LIMITS: these are temperatures that are incompatible with life. Lethal limits are established by
determining LD50s.

Lethal dose 50%: points of exposure where 50% of a population dies. Obviously this implies both the
temperature and duration of exposure to that temperature.

Why do different temperatures kill organisms? -- The effects of extreme temperatures

1. Cold -- the effects of freezing

a. physical damage to structures caused by the formation of ice: the membrane bound structures are
destroyed or damaged.
b. chemical damage due to the effects of high solute concentrations. When freezing occurs, solute
concentrations increase in non-frozen areas. These high concentrations may denature enzymes, etc.

2. Heat:

a. inadequate O2 supply for metabolic demands (especially in areas where O2 is low, such as water and
burrows)
b. rapid depletion of substrate stores

3. Heat and Cold

a. reduced activity or denaturation of proteins -- the inactivation of certain proteins with the result
that metabolic pathways are distorted.
b. disruption of enzyme pathways by differential temp effects on different enzymes (like b, except
what happens here is that temperature affects different reactions in different pathways differently.
c. effects on membrane fluidity:. These lead to problems with any membrane dependent function; the
nervous system is especially prone to disruption from fluidity changes. The types of fatty acids in a
membrane determine the motility of substances in the membrane and therefore their ability to interact
with each other and allow substances that require protein mediated transport to pass through the
membrane.
a. This effect is due to the inter-relationship between temperature (which causes a certain mean velocity
for membrane molecules) and the structure of the fatty acids making up the lipid bi-layer.
b. At a given temperature, the more saturated fatty acids that are present, the less fluid the membrane
will be. This is because saturated fatty acids are relatively straight chains and can easily be packed into a
membrane, producing a stable and not very fluid result.

By contrast, unsaturated fatty acids have bends at the location of every double bond; as a result,
they cannot pack as effectively into a membrane and the membrane tends to be more fluid.

GENERAL EFFECTS OF TEMPERATURE ON CHEMICAL AND PHYSICAL PROCESSES

1. CHEMICAL REACTION MEDIATED PROCESSES: all chemical reactions leading to such macro
phenomena as: muscle contractions, nerve transmission (both of these relate to sensation and
movement), digestion, growth, etc.

UNCATALYZED REACTIONS. Many chemical reactions commonly double their rate with a 10oC increase
in temperature.

2. PHYSICAL PROCESSES: these often dependent on temperature to a lesser degree than chemical
processes.

POIKILOTHERMY -- TEMPERATURE CONFORMITY

Many animals are totally incapable of temperature regulation. Their only significant source of heat is
the environment and their body temperature is directly determined by the ambient temperature. The
temperature conforming animals are referred to as poikilotherms. METABOLISM TENDS TO INCREASE
WITH BODY TEMPERATURE.

Poikilothermy is the condition of many organisms and all isolated tissues or cells. Many animals can
regulate their overall body temperature, their individual organs and tissues do not have this property
and act poikilothermically. Heart surgeons take advantage of this by perfusing the heart (and sometimes
other tissues also) with cool blood. This lowers the metabolism of the organ(s). Any organism can be
turned into a conformer -- when its ability to regulate is exceeded.

ECTOTHERMY: a term preferred to poikilotherm.

SUBHEADINGS of ectothermy, divided as to the identity of the major source of heat.

a. HELIOTHERM: the sun is the direct source of heat. So, these are ectotherms that use the sun as their
most significant direct heat source.

b. THIGMIOTHERM: warm substratum or medium (and not the sun directly) is the main heat source.

HOMEOTHERMY: this simply refers to a constant body temperature, presumably maintained by some
sort of regulatory mechanism. However, many also apply this term to animals that live in very constant
thermal environments that therefore have very little in the way of changes in body temperature.

ENDOTHERMY: the primary source of heat is internal chemical reactions.

HETEROTHERMY: An animal that acts like an endothermic homeotherm part of the time and like a
poikilothermic ectotherm the rest of the time.

The costs involved in temperature regulation fall primarily under two categories: TIME and ENERGY

a. Time is most important for ectotherms, to regulate temperature they must go in and out of the sun
or hide when cold.
b. Energy is the big cost for endotherms since the costs associated with a high rate of metabolism are so
large. It also involves a large time cost for many species due to the time they must spend looking for and
eating large amounts of food.

Both of these factors may result in great limitations on how small an animal can be and still be an
endotherm. They also relate to the particular ecological niche that is possible for an animal.

TEMPERATURE REGULATION: ENDOTHERMY*

I. Endothermic Temperature Regulation

Animals that more or less constantly regulate their body temperatures endothermically are often
referred to as being EUTHERMIC.

1. Eutherms include most all mammals and birds during most of their lives (and for many species, all of
their lives).

2. By contrast, another group of endotherms includes animals that only regulate their temperature
endothermically for a small portion of the time. These are referred to as HETEROTHERMS or
INTERMITTENT ENDOTHERMS. Many mammals and birds fall into these groups for at least part of their
lives. In addition, there are many species of insects and some fish and reptiles that also are intermittent
endotherms.

However, even in eutherms not all parts of the body are regulated at the same temperature. Some
definitions:

a. The core: the portions of the body, usually deep in the animal and most containing important organs
whose function is most dependent on a constant high body temperature. In most vertebrates the core
includes the brain and perhaps the heart and digestive system or the entire viscera.

b. The remainder of the animal is termed the periphery and is regulated to various degrees. For
instance, the temperature of the skin and to a lesser degree much of the limbs is less regulated than the
core. These areas may get relatively warm or even approach freezing with long-term harm or without
totally knocking out function.

ACCLIMATIZATION: the long-term (chronic) adjustments that are made to seasonal changes in
temperature.

ACCLIMATION: short-term changes quickly made to a changing set of conditions. This term is very often
applied to adjustments that are made by an animal to some specific set of laboratory conditions. Thus, it
is fair to say that they differ from acclimatization in being acute rather than chronic changes.
COMPENSATION: A process whose end result is to maintain a constant state regardless of the
conditions, i.e. a constant average daily metabolic rate regardless of the season through thermal
acclimatization.

Endothermy in Fish, Snakes, Turtles, And Lizards.

A. Fish

1. The difficulties that a potentially endothermic fish must face when compared to an endothermic
terrestrial animal:

a. The heat capacity of water is much higher than that of air and slightly higher than tissue (which is
mostly water). Thus, compared to a terrestrial animal, an aquatic animal experiences rapid loss of any
heat the animal generates.

b. O2 concentrations of water are much lower than in the air -- therefore to get a given amount of O2,
a fish must breathe a greater volume of water than must an air breather must breathe air:

In many fish there are two distinct groups of muscles: the largest mass is made up of white muscle
and is used for sprinting and the smaller mass is red muscle. The red muscle is located surrounding the
spine and the coelomic cavity (containing the gut and other organs). It is used for "cruising"; in other
words, it is continuously active as the fish makes its way around. The red muscle is the heat source:

a. it is continuously active
b. it is at the core whereas white muscle is nearer to the periphery
c. it surrounds the organs -- warming them will increase rates of digestion etc.

Other unusual examples of endothermy in vertebrates.

What body shape found in a terrestrial vertebrate would seem least conducive to endothermy?
Ans.: snake, due to large area of contact with the ground (much more thermally conductive than the
air).

At least one snake is an endotherm during part of its life: the Burmese Python, a common python
sold in many pet stores, it grows up to about 20 feet. The females wrap around their eggs and contract
their muscles to generate heat and incubate the eggs. During this time, the females maintain an
appreciably elevated body temperature.

Endothermy in Insects:

Flight is the most energetically demanding activity that any animal performs; there are many
situations where a high body temperature will aid in achieving flight.

1. This is most true in insects that have relatively great mass and small wings: i.e., high wing loading.
Examples are the highly maneuverable insect such as bees, dragonflies, some large flies, moths, and
beetles, and certain other insects such as some cicadas, crickets and grasshoppers.
2. Large insects with lightly loaded wings (e.g. butterflies, many moths) do not flap their wings often
enough to generate enough heat to be endothermic.
3. Many other insects are Behavioral Thermoregulators: this is especially true of butterflies.