Review session at 11:30 in INGR

2001
 Summary of things we skipped that I
still want you to know:
• About 7 L of fluid, mostly water, are absorbed in the gut daily.
• Most of the absorption occurs in the jejunum, and
• Water absorption involves a countercurrent exchange mechanism
that keeps [Na+] high in the tips of the villi.
• Hormones regulate gut processes
• Gastrin -- stimulates secretion of gastric juices and enhances
motility in several areas
• Secretin -- acts by several mechanisms to reduce acidity in the
duodenum
• Cholecystokinin (CCK) -- inhibits gastric motility and secretion,
stimulates secretion of pancreatic enzymes and release of bile, and
signals satiety
• Motilin -- stimulates motility in the stomach and small intestine
• Ghrelin -- stimulates growth hormone release and increases appetite
• Leptin – suppresses appetite
15.1 Introduction: Thermodynamics
and Life
• Life follows the laws of thermodynamics
• First Law of Thermodynamics
• Energy can neither be created nor destroyed.
• Energy is subject to input-output balance
in living systems.
• Law of the Conservation of Energy
15.1 Cont’d
• Second Law of Thermodynamics
• The entropy of a system and its surroundings
increases over time.
• Entropy is a measure of disorder in a system.
• The order a living system creates can arise
only at the expense of its environment.
• To compensate for entropy, animals require input of food
energy.
15.1 Cont’d
• Energy input-output balance
• Energy input
• Chemical energy is locked in the bonds of
food molecules and released when nutrient
molecules are broken down.
Energy input and output
Internal work is the energy-expending activities
occurring internally that maintain the body.
External work is the energy expended when
skeletal muscles are contracted to move the
body or to move external objects.
• Energy can change forms.
• Chemical energy of ATP is converted into
kinetic energy of locomotion.
• Almost all expended energy eventually
becomes heat.
• Energy is usually expressed in kilojoules (kJ) or
in units of heat (kilocalories or kcal)
• Example: 17.1 kJ or 4.1 kcal of heat energy is
released when 1 g of glucose is oxidized
15.2 Energy Balance:
General Principles
• The animal energy equation:

Einput – Eloss = ESMR/BMR + EActivity + EDIT + EProduction

• Input is the total food energy obtained.

• Loss is energy lost via feces, urine, etc.
• SMR and BMR are standard and basal metabolic rates -- the
minimum amount of energy needed to sustain waking life.
• Activity is the cost of neuromuscular efforts above the SMR or
BMR level.
• DIT is diet-induced thermogenesis -- the increase in metabolic
rate above basal level as a consequence of food intake.
• Production is the rate of energy storage.
Einput – Eloss = ESMR/BMR + EActivity + EDIT + EProduction

• Energy production determines

• net energy balance
• whether an animal will be at a stable
weight, lose weight, or gain weight
• Gaining significant mass is normal in
juvenile growth and in hibernators
laying down internal stores for winter,
but in other cases it leads to obesity
(BMI ≥ 30).
15.2 Cont’d
• Energy production cont’d
• States of balance
• Neutral energy balance -- occurs if energy input
equals energy output
• Positive energy balance -- if net energy intake
exceeds the amount of nonproduction energy
expended
• Negative energy balance -- If net energy intake
is less than the body’s immediate
nonproduction energy requirements
Einput – Eloss = ESMR/BMR + EActivity + EDIT + EProduction
• Metabolic rate is energy expenditure per unit time.
• Standard metabolic rate (SMR) is an
ectotherm’s resting metabolic rate at a
particular temperature (e.g. the average
temperature experienced naturally).
• Basal metabolic rate (BMR) is an endotherm’s
resting metabolic rate in its thermal neutral
zone.
Scaling: Metabolic rate as a function of
body mass
 Einput – Eloss = ESMR/BMR + EActivity + EDIT + EProduction

15.2 Cont’d
• Neuromuscular activity is potentially the
greatest energy usage for an animal.
• Activity includes locomotion powered by
skeletal muscles, breathing by respiratory
muscles and circulatory flow from the heart.
• Metabolic scope is the ratio of metabolic rate at
its highest level to the BMR or SMR.
• The metabolic cost of locomotion generally
increases with speed.
• More so with fish because of drag from a denser
medium.
Metabolic
rates (per
body mass)
for running
terrestrial
mammals
Metabolic
rate (per
whole
animal) for
a
swimming
fish
Metabolic
rate (per
body mass)
for flying
birds and
bats
Einput – Eloss = ESMR/BMR + EActivity + EDIT + EProduction
• Diet-induced thermogenesis (DIT)
occurs after eating.
• Obligatory DIT is the result of increased
metabolic activity associated with the
costs of obtaining and processing food.
• Regulatory DIT is increased heat
production after a meal for the purpose of
removing excess nutrients.
15.3 Energy Balance: Regulation
• Long-term regulation of energy balance
• Many adult mammals maintain fairly
constant mass over long periods.
• Homeostatic mechanisms maintain
long-term balance between energy
intake and output.
• Long-term regulation cont’d
• Periodic changes in body mass are adaptive in
other animals
• Hibernation
• Migration
• Pregnancy
• Many adult fish continue to grow larger if food
supplies permit
• Food intake is controlled by the
hypothalamus.
• Feeding and satiety centers are located in the
hypothalamus.
• Neuropeptide Y (NPY) is a potent appetite
stimulator.
• Melanocyte-stimulating hormone (α-MSH)
regulates skin color and suppresses appetite.
15.3 Cont’d
• Regulation by adipocytes
• Adipocytes secrete leptin in response to
increased fat storage.
• Leptin acts on the hypothalamus to inhibit NPY
secretion and stimulate α-MSH secretion.
• Leptin may be involved in the annual cycle of
hibernating mammals.
• Regulatory role of insulin
• Increased insulin secretion when
food is abundant inhibits NPY-
secreting cells of the
hypothalamus.
15.3 Cont’d
• Gastrointestinal hormones are involved in
initiation and cessation of eating.
• Cholecystokinin (CCK) signals satiety.
• Ghrelin stimulates NPY secretion.
• Levels rise before and at the onset of meals,
stimulating appetite.
• Obestatin signals satiety.
• PYY3-36 is the main antagonist of ghrelin.
• Inhibits NPY-secreting neurons
• Levels rise during meals, signaling satiety
 Melanocortins, e.g. MSH

Factors that
influence food
intake
15.3 Cont’d
• Effectors
• Regulation of eating is the most
important factor in the long-term
maintenance of energy balance and body
mass.
• Balance can be disrupted by psychosocial
and environmental influences (e.g. stress).
15.3 Cont’d
• Effectors cont’d
• Regulation of expenditures
• Underfeeding leads to energy-saving
reproductive adjustments (reducing fertility).
• Overfeeding leads to increased regulatory DIT.
• Increased activity of uncoupling proteins in
brown adipose tissue
• Increased activity after eating
• Both exercise and non-exercise activity
thermogenesis help dispose of excess
energy.
15.3 Cont’d
• Energy-balance set points have adapted
to availability of food supplies.
• Lean animals do well in habitats where food is
plentiful year-round.
• No need to store large amounts of body fat
• Lean animals are more agile than heavy
ones.
15.3 Cont’d
• Energy-balance set points cont’d
• A high adipose set point is valuable in habitats
where food is only abundant at limited times of
the year.
• Example: Weight gain before hibernation
• Hypothalami of these animals react only to
very high levels of leptin, allowing fat to
accumulate.
“Fat” mammals
15.4 Thermal Physiology:
General Principles
• Biological functions have an
optimal temperature range.
• Kinetic energy of reactants
increases with increasing
temperature.

Q10 = rate at temperature T

rate at temperature T-10
15.4 General Principles Cont’d
• Optimal temperature range
cont’d
• Denaturation of macromolecules
at temperatures above optimal.
• Some biomolecules can change
temperature optima over time
(acclimatization) or over many
generations (evolution)
The relative
activity of an
enzymatic
reaction as a
function of
temperature.
15.4 General Principles Cont’d
• Adaptive trade-offs between
thermostability and optimal
function.
• Cold-adapted animals have a higher
proportion of polyunsaturated fatty
acids in membranes, making the
membrane less viscous
(homeoviscous adaptation).
• Warm-adapted enzymes (e.g. lactate
dehydrogenase) are more rigid and
more resistant to denaturation
(homeoflexibility adaptation).
Saturation level of membrane
phospholipids changes with temperature
adaptation
Stiffening of
the
membrane
with
cholesterol.
Cholesterol
reduces
fluidity by
stabilizing
extended
forms of the
phospholipid
tails.
The relationship between adaptation temperature and ratio of
unsaturated phospholipid tails to saturated ones in gill
membranes of rainbow trout transferred to different temperatures
The relative
activity max
(Vmax) of
myosin ATPase
of four lizards
The enzyme
catalytic rate
constant (kcat)
for lactate
dehydrogenase
from various
vertebrates
adapted to
different
temperatures;
animals
adapted to
higher
temperatures
have enzymes
with lower
catalytic rates.
15.4 General Principles Cont’d
• Internal body temperature depends on
the difference between heat input and
heat output.
• Sources of heat input
• External environment (dominates in
ectotherms)
• Internal heat production (dominates in
endotherms)
• Sources of heat output
• Heat loss from exposed body surfaces
• Heat moves along a thermal gradient from
warmer to cooler regions.
Heat input and output
Mechanisms of heat transfer
15.4 General Principles Cont’d
• The body is a central core surrounded by an
outer shell.
• The core is made up of the abdominal and
thoracic organs, central nervous system and
skeletal muscles.
• Subject to regulation in animals that can
thermoregulate (whether ectotherm or
endotherm)
• The shell is made up of the skin and
subcutaneous fat.
• Temperature can vary and is usually cooler
than the core in endotherms.
15.4 Cont’d
• Balancing heat gain and heat loss to
regulate the core body temperature
• Gain external heat/avoid loss to cold
environs
• Heat gains are ectothermic sources made
by specific thermoregulatory behaviors
and aided by anatomic features.
• Retain internal heat
• Means include behavior, insulation,
reduced blood flow to the integument,
countercurrent exchangers and larger
body size.
15.4 General Principles Cont’d
• Balancing cont’d
• Generate more internal heat
• Heat-generating tissues in
endotherms
• Lose excess internal heat/avoid
gains from hot environs
• Mechanisms include behavior,
anatomy, increased blood flow to
the integument and increased
evaporation.
15.5 Ectothermy
• Ectotherms depend on external sources
for body heat.
• Most living things are poikilothermic.
• Poikilotherms’ body temperatures vary
with the environment.
• Thermoconformers have body
temperatures the same as the
environment (mostly aquatic animals, due
to high heat conductivity of water).
• Metabolic rates decrease in the cold.
• Avoid extreme temperatures; enter into
dormancy if unavoidable.
15.5 Ectothermy cont’d
•Ectothermic regulation
•An ectotherm in a
constant temperature
environment is
homeothermic.
15.5 Ectothermy cont’d
• Some ectotherms actively use external
heat exchanges to maintain their body
temperatures (e.g. lizard).
• Bask in the sun on cold mornings
• Constrict blood vessels in the skin to
reduce heat loss in a cold environment
• Dilate blood vessels in the skin to increase
heat loss in a warm environment
• A few ectotherms migrate to minimize
changes in body temperature (e.g.
monarch butterfly).
Relationship between body temperature
and air temperature for lizards
15.5 Ectothermy cont’d
• Metabolic compensation
• Animals’ metabolic reactions in cold
temperatures are increased to a level
close to that of warm-acclimated
animals.
• Increase level of metabolic enzymes
• Homeoviscous membrane adaptation
• Higher internal pH
• Isoform regulation -- different isoforms of
the same protein are used at different
temperatures
Activity of
lactate
dehydrogena
se in muscle
from
alligators
acclimatized
to winter and
summer
conditions
15.5 Ectothermy cont’d
• Survival of ectotherms in severe
cold
• Dormancy
• Turtles hibernate below the frost
line.
• Freeze tolerance
• Portion of body fluids freeze and
metabolic activity nearly stops
• Production of organic molecules
that serve as extracellular
antifreeze and intracellular
cryoprotectant osmolytes
15.5 Ectothermy cont’d
• Survival of ectotherms in severe
cold
• Freeze avoidance
• Use compatible cryoprotectants
throughout ECF and ICF (e.g.
antifreeze proteins)
• Supercooling -- maintaining liquid
water below the freezing point
• Cryoprotective dehydration --
increases osmolarity
15.5 Ectothermy cont’d
• Survival of ectotherms in extreme
heat
• Heat shock response
• A sudden increase in cellular
temperature causes heat shock factor-1
(HSF-1) trimers to activate genes for
heat shock proteins (HSPs).
• HSPs are small, thermostable,
hydophobic proteins that bind to larger
unfolded (denatured) proteins and assist
their folding into functional
conformations.
Model for regulation of hsp70, a major heat shock
protein whose activity is activated by sudden
temperature increases and other stresses
15.6 Endothermy and Homeothermy
• Endotherms depend mostly on
internal sources for body heat.
• Benefits of endothermy
• Less dependence on external heat
sources
• Faster biochemical processes
• Expansion of daily activity into nighttime
• Higher digestion rates
• Sustained locomotion
• Habitat expansion
• Regulated incubation temperature
15.6 Endothermy and Homeothermy cont’d

• Metabolic cost of
endothermy is large
• Endotherms must generate
heat internally.
• Must consume more energy
• Must have mechanisms to
remove excess heat
15.6 Endothermy and Homeothermy cont’d

• Birds and mammals are

homeotherms
• Homeotherms maintain a
consistent internal core
temperature.
• Rely on high levels of aerobic
metabolism for sustainable
heat production
15.6 Endothermy and Homeothermy cont’d

• Core body temperatures vary

among species of birds and
mammals
• Birds have higher set points than
mammals.
• Placental mammals have higher set
points than monotremes.
• Body temperatures vary among
individuals and are lower at night
than during the day.
15.6 Endothermy and Homeothermy cont’d

 Balancing heat gain and heat

loss to regulate the core body
temperature
• Gain external heat/avoid loss to
cold environs
• Ectothermic behavior (e.g.
basking in sun)
• Anatomic features that help absorb
radiation (e.g. dark skin or
feathers)
 A
roadrunner
ruffles its
feathers to
expose dark
skin, which
absorbs
sunlight to
help heat
the animal’s
blood.
15.6 Endothermy and Homeothermy cont’d
 Balancing heat gain and loss cont’d
• Retain internal heat
• Reduced blood flow to integument through
vasoconstriction
• Anatomic insulation (feathers, hair,
subcutaneous adipose layers) and piloerection
• Behavioral insulation (nesting, burrowing,
postural changes, huddling)
• Larger body size in colder climate
• Countercurrent exchangers (rete mirabile in the
limbs, maxilloturbinal folds in the nasal cavity)
Changes in blood flow to the skin for thermoregulation
Countercurrent
heat
exchangers
(retes) in
endotherms
15.6 Endothermy and Homeothermy cont’d

 Balancing heat gain and loss

cont’d
• Generate more internal heat
• BMR is 5-20 times greater than
SMR of an ectotherm of the same
mass.
• Increased muscular activity and
shivering
• Nonshivering thermogenesis (e.g.
brown adipose tissue)
Brown
adipose
tissue (BAT)
in
endotherms
15.6 Endothermy and Homeothermy cont’d
 Balancing heat gain and loss cont’d
• Lose excess internal heat/avoid gains from
hot environs
• Reduced insulation
• Increased blood flow to integument through
vasodilation
• Enhanced evaporation (panting, insensible
cutaneous loss, sweating)
• Countercurrent exchange (rete protecting brain
from overheating)
• Avoidance behavior (seeking shade or seasonal
migration)
• Anatomic features that reflect radiation (e.g. light
skin or feathers)
15.6 Endothermy and Homeothermy cont’d
• Thermostat is in the hypothalamus.
• Birds also have a reflex integrator in the spinal cord.

• Central thermoreceptors
monitor core temperature.
• Peripheral thermoreceptors
monitor skin temperature to
provide anticipatory
information.
15.6 Endothermy and Homeothermy cont’d

• Thermostat cont’d
• Posterior region of hypothalamus
• Activated by cold
• Triggers reflexes that mediate heat
production and heat conservation
• Anterior region of hypothalamus
• Activated by warmth
• Initiates reflexes that mediate heat
loss
Major thermoregulatory pathways in a mammal
15.6 Endothermy and Homeothermy cont’d
• Coordinated regulation of body
temperature
• The thermal neutral zone (TNZ) is a
range of environmental temperatures
in which the animal does not need to
expend significant energy for
thermoregulation.
• At the lower critical temperature,
metabolic rate increases as special
heat-generating mechanisms are
activated
• Hypothermia is a dangerously low body
temperature.
15.6 Endothermy and Homeothermy cont’d
• Coordinated regulation cont’d
• At the upper critical temperature,
metabolic rate increases due to
panting or heavy sweating.
• Hyperthermia is a dangerously high
body temperature.
• Fever is an elevation in body
temperature as a result of an infection
or inflammation.
• The hypothalamic thermostat is reset to
a higher set point.
The influence of ambient temperature on the
metabolic rate (MR) of a small temperate
endotherm, showing the thermoneutral zone
where MR is constant.
15.7 Heterothermy
• Heterotherms are animals that have
endothermy, but are not fully
homeotherms.
• Regional heterothermy
• Endothermic animals that heat only
certain organs or body regions.
• Laminid sharks and scombroid teleosts
have endothermic aerobic swimming
muscles; countercurrent rete system
between aerobic muscle and gills
reduces heat loss from the gills
Countercurrent exchangers in heterothermic tuna.
The diagram shows the rete located between the outer
(cutaneous) vessels and the deep red muscle. The rete
traps the heat produced by those muscles.
15.7 Heterothermy cont’d
• Heterotherms are animals that have
endothermy but are not fully homeotherms

• Temporal heterothermy
•Endothermic animals that
regularly shift from a
regulated high body
temperature to a low
body temperature.
15.7 Heterothermy cont’d
• Torpor is a short-term dormant
state accompanied by a drop in
body temperature.
• Small endotherms with high
metabolic demands (e.g. shrews
and birds) enter torpor on a daily
basis.
• Torpor at night conserves
energy.
• Animals that employ daily torpor
have longer life spans.
15.7 Heterothermy cont’d
• Temporal heterothermy cont’d
• Hibernation is a long-term dormant
state in which body temperature
drops to near ambient temperature.
• Hibernators store up large amounts of
unsaturated fats to serve as energy
reserves.
• Have a functioning thermostat with a
greatly lowered set point
• Only small mammals hibernate; bears
undergo winter sleep in which body
temperature drops about 5°C.
Temperatures
of mammals
in torpor and
hibernation
15.8 Thermal Physiology and
Climate Change
• Global warming is due in part to
anthropogenic causes.
• Increasing levels of greenhouse gases
(mainly CO2) from burning fossil fuels
prevent infrared radiation from leaving
the Earth.
• The Arctic region has experienced the
greatest warming.
• Climate change is expected to impact
all types of organisms.
Global temperature
changes and their effects
on animal metabolic rates
15.8 Thermal Physiology and
Climate Change
• Observed impacts of climate
change on animals
• Heat stress as severe heat waves have
increased
• Energy balance is disrupted by warming
environments resulting in large-scale
deaths
• Habitat change - populations are shifting
northward (in the Northern Hemisphere)
or to higher altitudes
• Reproduction - earlier breeding seasons