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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.