Fluid dynamics: hydrodynamics

of water and other fluids, fluid

dynamics of
blood
Hydrodynamics of
Water
Water as continuum: Though Fluids Are Composed of Molecules It Is
Possible to Treat Them as a Continuous Medium
Overview: The Molecule That Supports
All of Life
• Water is the biological medium on Earth
• All living organisms require water more than any other
substance
• Most cells are surrounded by water, and cells
themselves are about 70–95% water
• The abundance of water is the main reason the Earth is
habitable

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Fig. 3-1
Concept 3.1: The polarity of water
molecules results in hydrogen bonding
• The water molecule is a polar molecule: The opposite
ends have opposite charges
• Polarity allows water molecules to form hydrogen
bonds with each other

Animation: Water Structure

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Fig. 3-2

–
Hydrogen
+ bond
H

——
O
–
+
——
H
– +
–
+
Fig. 3-UN1
Concept 3.2: Four emergent properties
of water contribute to Earth’s fitness for
life
• Four of water’s properties that facilitate an
environment for life are:
• Cohesive behavior
• Ability to moderate temperature
• Expansion upon freezing
• Versatility as a solvent

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Cohesion
• Collectively, hydrogen bonds hold water molecules
together, a phenomenon called cohesion
• Cohesion helps the transport of water against gravity in
plants
• Adhesion is an attraction between different substances,
for example, between water and plant cell walls

Animation: Water Transport

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Fig. 3-3

Adhesion

Water-conducting
cells

Direction Cohesion
of water
150 µm
movement
• Surface tension is a measure of how hard it is to break
the surface of a liquid
• Surface tension is related to cohesion

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Fig. 3-4
Moderation of Temperature

• Water absorbs heat from warmer air and releases

stored heat to cooler air
• Water can absorb or release a large amount of heat
with only a slight change in its own temperature

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Heat and Temperature

• Kinetic energy is the energy of motion

• Heat is a measure of the total amount of kinetic energy
due to molecular motion
• Temperature measures the intensity of heat due to the
average kinetic energy of molecules

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• The Celsius scale is a measure of temperature using
Celsius degrees (°C)
• A calorie (cal) is the amount of heat required to raise
the temperature of 1 g of water by 1°C
• The “calories” on food packages are actually
kilocalories (kcal), where 1 kcal = 1,000 cal
• The joule (J) is another unit of energy where
1 J = 0.239 cal, or 1 cal = 4.184 J

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Water’s High Specific Heat

• The specific heat of a substance is the amount of heat

that must be absorbed or lost for 1 g of that substance
to change its temperature by 1ºC
• The specific heat of water is 1 cal/g/ºC
• Water resists changing its temperature because of its
high specific heat

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• Water’s high specific heat can be traced to hydrogen
bonding
• Heat is absorbed when hydrogen bonds break
• Heat is released when hydrogen bonds form
• The high specific heat of water minimizes temperature
fluctuations to within limits that permit life

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Fig. 3-5

Burbank San Bernardino

Santa Barbara 73°
90° 100°
Los Angeles Riverside 96°
(Airport) 75° Santa Ana
Palm Springs
70s (°F) 84°
106°
80s Pacific Ocean
90s
100s San Diego 72°

40 miles
Evaporative Cooling

• Evaporation is transformation of a substance from

liquid to gas
• Heat of vaporization is the heat a liquid must absorb for
1 g to be converted to gas
• As a liquid evaporates, its remaining surface cools, a
process called evaporative cooling
• Evaporative cooling of water helps stabilize
temperatures in organisms and bodies of water

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Insulation of Bodies of Water by
Floating Ice
• Ice floats in liquid water because hydrogen bonds in ice
are more “ordered,” making ice less dense
• Water reaches its greatest density at 4°C
• If ice sank, all bodies of water would eventually freeze
solid, making life impossible on Earth

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 3-6

Hydrogen
bond
Ice Liquid water
Hydrogen bonds are stable Hydrogen bonds break and re-form
Fig. 3-6a

Hydrogen
bond
Ice Liquid water
Hydrogen bonds are stable Hydrogen bonds break and re-form
The Solvent of Life

• A solution is a liquid that is a homogeneous mixture of

substances
• A solvent is the dissolving agent of a solution
• The solute is the substance that is dissolved
• An aqueous solution is one in which water is the
solvent

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• Water is a versatile solvent due to its polarity, which
allows it to form hydrogen bonds easily
• When an ionic compound is dissolved in water, each ion
is surrounded by a sphere of water molecules called a
hydration shell

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 3-7

–
Na+
+
– –+
+
– –
Na+ –
+ +

Cl– Cl– + –
–
+
–
+
–
–
• Water can also dissolve compounds made of nonionic
polar molecules
• Even large polar molecules such as proteins can dissolve
in water if they have ionic and polar regions

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 3-8

(a) Lysozyme molecule in a (b) Lysozyme molecule (purple) in an aqueous (c) Ionic and polar regions
nonaqueous environment environment on the protein’s surface
attract water molecules.
Fig. 3-8ab

(a) Lysozyme molecule in a (b) Lysozyme molecule (purple) in an aqueous

nonaqueous environment environment
Fig. 3-8bc

(b) Lysozyme molecule (purple) in an aqueous (c) Ionic and polar regions
environment on the protein’s surface
attract water molecules.
Hydrophilic and Hydrophobic
Substances
• A hydrophilic substance is one that has an affinity for
water
• A hydrophobic substance is one that does not have an
affinity for water
• Oil molecules are hydrophobic because they have
relatively nonpolar bonds
• A colloid is a stable suspension of fine particles in a
liquid

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Solute Concentration in Aqueous
Solutions
• Most biochemical reactions occur in water
• Chemical reactions depend on collisions of molecules
and therefore on the concentration of solutes in an
aqueous solution

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• Molecular mass is the sum of all masses of all atoms in
a molecule
• Numbers of molecules are usually measured in moles,
where 1 mole (mol) = 6.02 x 1023 molecules
• Avogadro’s number and the unit dalton were defined
such that 6.02 x 1023 daltons = 1 g
• Molarity (M) is the number of moles of solute per liter
of solution

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Concept 3.3: Acidic and basic
conditions affect living organisms
• A hydrogen atom in a hydrogen bond between two
water molecules can shift from one to the other:
• The hydrogen atom leaves its electron behind and is
transferred as a proton, or hydrogen ion (H+)
• The molecule with the extra proton is now a hydronium ion
(H3O+), though it is often represented as H+
• The molecule that lost the proton is now a hydroxide ion
(OH–)

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• Water is in a state of dynamic equilibrium in which
water molecules dissociate at the same rate at which
they are being reformed

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 3-UN2

H
H
O H O O H O
H H H H

2H2O Hydronium Hydroxide

ion (H3O+) ion (OH–)
• Though statistically rare, the dissociation of water
molecules has a great effect on organisms
• Changes in concentrations of H+ and OH– can drastically
affect the chemistry of a cell

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Effects of Changes in pH

• Concentrations of H+ and OH– are equal in pure water

• Adding certain solutes, called acids and bases, modifies
the concentrations of H+ and OH–
• Biologists use something called the pH scale to describe
whether a solution is acidic or basic (the opposite of
acidic)

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Acids and Bases

• An acid is any substance that increases the H+

concentration of a solution
• A base is any substance that reduces the H+
concentration of a solution

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The pH Scale

• In any aqueous solution at 25°C the product of H+ and

OH– is constant and can be written as
[H+][OH–] = 10–14
• The pH of a solution is defined by the negative
logarithm of H+ concentration, written as
pH = –log [H+]
• For a neutral aqueous solution
[H+] is 10–7 = –(–7) = 7

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• Acidic solutions have pH values less than 7
• Basic solutions have pH values greater than 7
• Most biological fluids have pH values in the range of 6
to 8

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Fig. 3-9
pH Scale
0

1
Battery acid
Gastric juice,
2 lemon juice

Increasingly Acidic
H+
H+
H+
3 Vinegar, beer,
–
H+ OH
+
OH– H H+ wine, cola

[H+] > [OH–]

H+ H+

Acidic 4 Tomato juice

solution
Black coffee
5
Rainwater
6 Urine

OH–
OH– Saliva
Neutral
H +
H + OH– 7 Pure water
[H+] = [OH–]
OH– OH– + Human blood, tears
H+ H+ H
8 Seawater
Neutral
solution
9

Increasingly Basic
10

[H+] < [OH–]

Milk of magnesia
OH–
OH–
11
OH– H+ OH–
OH OH
– – Household ammonia
H+ OH–
12
Basic
solution Household
13 bleach
Oven cleaner
14
Buffers

• The internal pH of most living cells must remain close to

pH 7
• Buffers are substances that minimize changes in
concentrations of H+ and OH– in a solution
• Most buffers consist of an acid-base pair that reversibly
combines with H+

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Threats to Water Quality on Earth
• Acid precipitation refers to rain, snow, or fog with a pH
lower than 5.6
• Acid precipitation is caused mainly by the mixing of
different pollutants with water in the air and can fall at
some distance from the source of pollutants
• Acid precipitation can damage life in lakes and streams
• Effects of acid precipitation on soil chemistry are
contributing to the decline of some forests

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Fig. 3-10

0 More
1 acidic
2
3 Acid
4 rain
5
Normal
6 rain
7
8
9
10
11
12
13 More
14 basic
• Human activities such as burning fossil fuels threaten
water quality
• CO2 is released by fossil fuel combustion and contributes
to:
• A warming of earth called the “greenhouse” effect
• Acidification of the oceans; this leads to a decrease in the
ability of corals to form calcified reefs

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 3-11
EXPERIMENT

RESULTS

40
Calcification rate

per m2 per day)

(mmol CaCO3

20

0
150 200 250 300
[CO32–] (µmol/kg)
Fig. 3-11a

EXPERIMENT
Fig. 3-11b

RESULTS

Calcification rate

per m2 per day)

40
(mmol CaCO3
20

0
150 200 250 300
[CO32–] (µmol/kg)
Fig. 3-UN3

–
Hydrogen
+ bond
H

– O
+ H
– +
+ –
Fig. 3-UN4

Ice: stable hydro- Liquid water:

gen bonds transient hydrogen
bonds
Fig. 3-UN5
0
Acidic
[H+] > [OH–]
Acids donate H+ in
aqueous solutions

Neutral
[H+] = [OH–] 7

Bases donate OH–

or accept H+ in
Basic aqueous solutions
[H+] < [OH–]
14
Fig. 3-UN6

Surface of Mars Surface of Earth

Fig. 3-UN7
You should now be able to:

1. List and explain the four properties of water that

emerge as a result of its ability to form hydrogen
bonds
2. Distinguish between the following sets of terms:
hydrophobic and hydrophilic substances; a solute, a
solvent, and a solution
3. Define acid, base, and pH
4. Explain how buffers work

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Fluid Dynamics of
Blood
Overview: Trading Places

• Every organism must exchange materials with its

environment.
• Exchanges ultimately occur at the cellular level.
• In unicellular organisms, these exchanges occur
directly with the environment.
• For most cells making up multicellular organisms,
direct exchange with the environment is not
possible.
• Gills are an example of a specialized exchange
system in animals.
• Internal transport and gas exchange are functionally
related in most animals.
How does a feathery fringe help this animal survive?
Circulatory systems link exchange
surfaces with cells throughout the body
• In small and/or thin animals, cells can exchange
materials directly with the surrounding medium.
• In most animals, transport systems connect the
organs of exchange with the body cells.
• Most complex animals have internal transport
systems that circulate fluid.
Gastrovascular Cavities
• Simple animals, such as cnidarians, have a body wall
that is only two cells thick and that encloses a
gastrovascular cavity.
• This cavity functions in both digestion and
distribution of substances throughout the body.
• Some cnidarians, such as jellies, have elaborate
gastrovascular cavities.
• Flatworms have a gastrovascular cavity and a large
surface area to volume ratio.
Internal transport in gastrovascular cavities

Circular
canal

Mouth
Pharynx
Mouth
Radial canal 5 cm 2 mm

(a) The moon jelly Aurelia, a cnidarian (b) The planarian Dugesia, a
flatworm
Open and Closed Circulatory Systems

• More complex animals have either open or closed

circulatory systems.
• Both systems have three basic components:
• A circulatory fluid = blood or hemolymph.
• A set of tubes = blood vessels.
• A muscular pump = the heart.
• In insects, other arthropods, and most molluscs,
blood bathes the organs directly in an open
circulatory system.
• In an open circulatory system, there is no distinction
between blood and interstitial fluid, and this general
body fluid is more correctly called hemolymph.
• In a closed circulatory system, the blood is confined
to vessels and is distinct from the interstitial fluid.
• Closed systems are more efficient at transporting
circulatory fluids to tissues and cells.
Open and closed circulatory systems

Heart Heart

Blood
Hemolymph in
sinuses Interstitial Small branch vessels
surrounding organs fluid In each organ

Pores
Dorsal vessel
(main heart)

Tubular heart Auxiliary hearts Ventral vessels

(a) An open circulatory system (b) A closed circulatory system
Organization of Vertebrate Closed
Circulatory Systems
• Humans and other vertebrates have a closed
circulatory system, often called the cardiovascular
system.
• The three main types of blood vessels are:
arteries - away from the heart.
veins - toward the heart.
capillaries - exchange with body cells.
• Arteries branch into arterioles and carry blood to
capillaries.
• Networks of capillaries called capillary beds are the
sites of chemical exchange between the blood and
interstitial fluid.
• Venules converge into veins and return blood from
capillaries to the heart.
• Vertebrate hearts contain two or more chambers.
• Blood enters through an atrium and is pumped out
through a ventricle.
Atria - receive blood
Ventricles - pump blood
Single Circulation

• Bony fishes, rays, and sharks have single circulation

with a two-chambered heart.
• In single circulation, blood leaving the heart passes
through two capillary beds before returning.
Single circulation in fishes

Gill capillaries

Artery Gill
circulation

Ventricle
Heart
Atrium

Systemic
Vein circulation

Systemic capillaries
Double Circulation

• Amphibian, reptiles, and mammals have double

circulation.
• Oxygen-poor and oxygen-rich blood are pumped
separately from the right and left sides of the heart.
Double circulation in vertebrates

Mammals and
Amphibians Reptiles Birds

Lung and skin capillaries Lung capillaries Lung capillaries

Pulmocutaneous Right
Pulmonary Pulmonary
circuit systemic circuit circuit
aorta

Atrium (A) Atrium (A) A A A A

Ventricle (V) V V Left V V
Right Left Right Left systemic Right Left
Systemic aorta Systemic
circuit circuit

Systemic capillaries Systemic capillaries Systemic capillaries

• In reptiles and mammals, oxygen-poor blood flows
through the pulmonary circuit to pick up oxygen
through the lungs.
• In amphibians, oxygen-poor blood flows through a
pulmocutaneous circuit to pick up oxygen through
the lungs and skin.
• Oxygen-rich blood delivers oxygen through the
systemic circuit.
• Double circulation maintains higher blood pressure
in the organs than does single circulation.
Adaptations of Double Circulatory
Systems
Amphibians:
• Frogs / amphibians have a three-chambered heart:
2 atria and 1 ventricle.
• The ventricle pumps blood into a forked artery that
splits the ventricle’s output into the
pulmocutaneous circuit and the systemic circuit.
• Underwater, blood flow to the lungs is nearly shut
off.
Reptiles (Except Birds)
• Turtles, snakes, and lizards have a three-chambered
heart: two atria and one ventricle.
• In alligators, caimans, and other crocodilians a
septum - partially or fully divides the ventricle.
• Reptiles have double circulation, with a pulmonary
circuit - lungs and a systemic circuit.
 RA --> RV --> LUNGS --> LA --> LV --> Body

Mammals
• Mammals and birds have a four-chambered heart
with two atria and two ventricles.
• The left side of the heart pumps and receives only
oxygen-rich blood, while the right side receives and
pumps only oxygen-poor blood.
• Mammals and birds are endotherms and require
more O2 than ectotherms.
Coordinated cycles of heart contraction
drive double circulation in mammals
• Blood begins its flow with the right ventricle
pumping blood to the lungs.
• In the lungs, the blood loads O2 and unloads CO2
• Oxygen-rich blood from the lungs enters the heart
at the left atrium and is pumped through the aorta
to the body tissues by the left ventricle.
• The aorta provides blood to the heart through the
coronary arteries.
• Blood returns to the heart through the superior
vena cava (deoxygenated blood from head, neck,
and forelimbs) and inferior vena cava
(deoxygenated blood from trunk and hind limbs).
• The superior vena cava and inferior vena cava flow
into the Right Atrium - RA.
 Superior vena cava 7
Capillaries of head and
Returns deoxygenated blood from Forelimbs - EXCHANGE
body to heart RA

Pulmonary artery Pulmonary artery

Carries deoxygenated blood to lungs

Capillaries Aorta
9 Capillaries
of right Lung
GAS EXCHANGE of left Lung
GAS EXCHANGE

3 2 3
4
11
Pulmonary vein
Pulmonary vein Carries oxygenated blood
5 to heart: LA
1
Right Atrium Left Atrium - LA
RA - Receives deoxygenated blood 10 Receives oxygenated blood
from body from lungs
Right Ventricle Left Ventricle - LV
RV - Pumps blood to lungs Pumps oxygenated blood to body
Inferior vena cava Aorta = main artery to body
Returns deoxygenated blood from
mammalian cardiovascular system
for Systemic Circulation
body to heart RA
Capillaries of
8 abdominal organs and hind limbs
EXCHANGE with body cells
The Mammalian Heart: A Closer Look

• A closer look at the mammalian heart provides a

better understanding of double circulation.
• RIGHT side = deoxygenated blood from body
pumped to lungs.
• LUNGS = gas exchange.
• LEFT side = oxygenated blood from lungs pumped to
body.
 Mammalian Heart
Pulmonary artery Aorta - systemic
- to lungs circulation

Right Atrium RA Pulmonary veins -

Receives from lungs to heart
Deoxygented
Blood from Left Atrium LA
body Receives oxgenated
blood from lungs

Semilunar Semilunar
valve valve

Atrioventricular Atrioventricular
valve valve

Right Ventricle RV Left Ventricle LV

Pumps to lungs for Pumps oxygenated
gas exchange blood to body via aorta
• The heart contracts and relaxes in a rhythmic cycle
called the cardiac cycle.
• The contraction, or pumping, phase is called
systole.
• The relaxation, or filling, phase is called diastole.
• Blood Pressure = systolic / diastolic
Cardiac cycle
2 Atrial systole;
Semilunar ventricular
valves diastole
closed

0.1 sec

Semilunar
AV valves
valves 0.4 sec 0.3 sec open
open

1 Atrial and
ventricular
diastole
AV valves
closed
3 Ventricular systole;
atrial diastole
• The heart rate, also called the pulse, is the number
of beats per minute.
• The stroke volume is the amount of blood pumped
in a single contraction.
• The cardiac output is the volume of blood pumped
into the systemic circulation per minute and
depends on both the heart rate and stroke volume.
Four valves prevent backflow of blood in the heart:
• The atrioventricular (AV) valves separate each atrium and
ventricle.
• The semilunar valves control blood flow to the aorta and
the pulmonary artery.
• The “lub-dup” sound of a heart beat is caused by the recoil
of blood against the AV valves (lub) then against the
semilunar (dup) valves.
• Backflow of blood through a defective valve causes a heart
murmur.
Maintaining the Heart’s Rhythmic Beat
• Some cardiac muscle cells are self-excitable = they contract
without any signal from the nervous system.
• The sinoatrial (SA) node, or pacemaker, sets the rate and
timing at which cardiac muscle cells contract.
• Impulses from the SA node travel to the atrioventricular
(AV) node. At the AV node, the impulses are delayed and
then travel to the Purkinje fibers that make the ventricles
contract.
• Impulses that travel during the cardiac cycle can be
recorded as an electrocardiogram (ECG or EKG). The
pacemaker is influenced by nerves, hormones, body
temperature, and exercise.
Control of heart rhythm

1 Pacemaker 2 Signals are 3 Signals pass 4 Signals spread

generates wave of delayed at to heart apex. throughout
signals to contract. AV node. ventricles.

AV
SA node node
(pacemaker) Bundle Purkinje Fibers:
branches Heart
ventricles contract
apex

ECG
Patterns of blood pressure and flow
reflect the structure and arrangement of
blood vessels
• The physical principles that govern movement of
water in plumbing systems also influence the
functioning of animal circulatory systems.
• The epithelial layer that lines blood vessels is called
the endothelium.
Structure
of
blood vessels

Artery Vein

SEM
100 µm Valve
Basal lamina

Endothelium Endothelium
Smooth Smooth
muscle muscle
Connective Connective
tissue Capillary tissue
Artery Vein

Arteriole Venule

15 µm
Red blood cell

Capillary

LM
• Capillaries have thin walls, the endothelium plus its
basement membrane, to facilitate the exchange of
materials.
• Arteries and veins have an endothelium, smooth
muscle, and connective tissue.
• Arteries have thicker walls than veins to
accommodate the high pressure of blood pumped
from the heart.
• In the thinner-walled veins, blood flows back to the
heart mainly as a result of muscle action.
Blood Flow Velocity

• Physical laws governing movement of fluids through

pipes affect blood flow and blood pressure.
• Velocity of blood flow is slowest in the capillary
beds, as a result of the high resistance and large
total cross-sectional area.
• Blood flow in capillaries is necessarily slow for
exchange of materials.
The interrelationship of cross-sectional area of blood vessels,
blood flow velocity, and
blood pressure.

5,000

Area (cm2)
4,000
3,000
2,000
1,000
0

50
40

(cm/sec)
Velocity
30
20
10
0
120
Systolic
100
Pressure

pressure
(mm Hg)

80
60
40 Diastolic
20 pressure
0

Capillaries
Venules
Arterioles
Aorta

Veins

Venae cavae
Arteries
Blood Pressure

• Blood pressure is the hydrostatic pressure that

blood exerts against the wall of a vessel.
• In rigid vessels blood pressure is maintained; less
rigid vessels deform and blood pressure is lost.
Changes in Blood Pressure During the
Cardiac Cycle
• Systolic pressure is the pressure in the arteries
during ventricle contraction /systole; it is the
highest pressure in the arteries.
• Diastolic pressure is the pressure in the arteries
during relaxation /diastole; it is lower than systolic
pressure.
• A pulse is the rhythmic bulging of artery walls with
each heartbeat.
Regulation of Blood Pressure

• Blood pressure is determined by cardiac output and

peripheral resistance due to constriction of
arterioles.
• Vasoconstriction is the contraction of smooth
muscle in arteriole walls; it increases blood
pressure.
• Vasodilation is the relaxation of smooth muscles in
the arterioles; it causes blood pressure to fall.
• Vasoconstriction and vasodilation help maintain adequate
blood flow as the body’s demands change.
• The peptide endothelin is an important inducer of
vasoconstriction.
• Blood pressure is generally measured for an artery in the
arm at the same height as the heart.
• Blood pressure for a healthy 20 year old at rest is 120 mm
Hg at systole / 70 mm Hg at diastole.
Question: How do endothelial cells control vasoconstriction?

RESULTS

Ser
Leu
Ser Endothelin
Met Cys Ser Cys —NH +
3
Asp
Lys
Glu Cys Val Tyr Phe Cys His Leu Asp Ile Ile Trp —COO–

Cys Trp
Parent polypeptide
1 53 73

Endothelin 203
Measurement of blood pressure:
sphygmomanometer

Blood pressure reading: 120/70

Pressure in cuff Pressure in cuff Pressure in
greater than drops below cuff below
120 mm Hg 120 mm Hg 70 mm Hg

Rubber
cuff
inflated
120 120
with air
70

Artery Sounds Sounds

closed audible in stop
stethoscope
• Fainting is caused by inadequate blood flow to the
head.
• Animals with longer necks require a higher systolic
pressure to pump blood a greater distance against
gravity.
• Blood is moved through veins by smooth muscle
contraction, skeletal muscle contraction, and
expansion of the vena cava with inhalation.
• One-way valves in veins / heart prevent backflow
of blood.
Blood flow in veins

Direction of blood flow

in vein (toward heart)
Valve (open)

Skeletal muscle

Valve (closed)
Capillary Function

• Capillaries in major organs are usually filled to

capacity. Blood supply varies in many other sites.
• Two mechanisms regulate distribution of blood in
capillary beds:
• Contraction of the smooth muscle layer in the wall of an
arteriole constricts the vessel.
• Precapillary sphincters control flow of blood between
arterioles and venules.
Blood flow in capillary beds

Precapillary sphincters Thoroughfare

channel

Capillaries
Arteriole Venule

(a) Sphincters relaxed

Arteriole Venule

(b) Sphincters contracted

• The critical exchange of substances between the
blood and interstitial fluid takes place across the
thin endothelial walls of the capillaries.
• The difference between blood pressure and
osmotic pressure drives fluids out of capillaries at
the arteriole end and into capillaries at the venule
end.
Fluid exchange between capillaries and the interstitial fluid

Body tissue
INTERSTITIAL FLUID
Capillary
Net fluid
movement out
Net fluid
movement in

Direction of
blood flow

Blood pressure = hydrostatic pressure

Pressure

Inward flow

Outward flow
Osmotic pressure

Arterial end of capillary Venous end

Fluid Return by the Lymphatic System

• The lymphatic system - returns fluid that leaks out

in the capillary beds … restoring filtered fluid to
blood maintains homeostasis.
• This system aids in body defense.
• Fluid, called lymph, reenters the circulation directly
at the venous end of the capillary bed and indirectly
through the lymphatic system.
• The lymphatic system drains into neck veins.
• Lymph nodes are organs that produce phagocytic
white blood cells and filter lymph - an important
role in the body’s defense.
• Edema is swelling caused by disruptions in the flow
of lymph.
Blood Composition and Function

• Blood consists of several kinds of blood cells

suspended in a liquid matrix called plasma.
• The cellular elements: red blood cells, white blood
cells, and platelets occupy about 45% of the volume
of blood.
Composition of mammalian blood

Plasma 55%
Constituent Major functions Cellular elements 45%
Cell type Number Functions
Water Solvent for
per µL (mm3) of blood
carrying other
substances
Erythrocytes 5–6 million Transport oxygen
(red blood cells) and help transport
Ions (blood electrolytes) carbon dioxide
Sodium Osmotic balance, Separated
Potassium pH buffering, and blood
Calcium regulation of elements
Magnesium membrane
Chloride permeability
Bicarbonate Leukocytes 5,000–10,000 Defense and
(white blood cells) immunity
Plasma proteins
Albumin Osmotic balance
pH buffering
Basophil Lymphocyte
Fibrinogen Clotting
Immunoglobulins Defense Eosinophil
(antibodies)

Neutrophil Monocyte
Substances transported by blood
Nutrients (such as glucose, fatty acids, vitamins)
Waste products of metabolism Platelets 250,000– Blood clotting
Respiratory gases (O2 and CO2) 400,000
Hormones
Plasma
• Blood plasma is about 90% water.
• Among its solutes are inorganic salts in the form of
dissolved ions, sometimes called electrolytes.
• Another important class of solutes is the plasma
proteins, which influence blood pH, osmotic
pressure, and viscosity. Various plasma proteins
function in lipid transport, immunity, and blood
clotting.
• Plasma transports nutrients, gases, and cell waste.
Cellular Elements

• Suspended in blood plasma are two types of cells:

• Red blood cells rbc = erythrocytes, transport oxygen.
• White blood cells wbc = leukocytes, function in defense.
• Platelets are fragments of cells that are involved in
blood clotting.
Erythrocytes - Oxygen Transport
• Red blood cells, or erythrocytes, are by far the most
numerous blood cells.
• They transport oxygen throughout the body.
• They contain hemoglobin, the iron-containing
protein that transports oxygen.
Leukocytes - Defense
• There are five major types of white blood cells, or
leukocytes: monocytes, neutrophils, basophils,
eosinophils, and lymphocytes.
• They function in defense by phagocytizing bacteria
and debris or by producing antibodies.
• They are found both in and outside of the
circulatory system.
 Platelets - Blood Clotting
• Platelets are fragments of cells and function in
blood clotting.
• When the endothelium of a blood vessel is
damaged, the clotting mechanism begins.
• A cascade of complex reactions converts fibrinogen
to fibrin, forming a clot.
• A blood clot formed within a blood vessel is called a
thrombus and can block blood flow.
Blood clotting

Red blood cell

Collagen fibers
Platelet plug Fibrin clot
Platelet releases chemicals
that make nearby platelets sticky
Clotting factors from:
Platelets
Damaged cells
Plasma (factors include calcium, vitamin K)

Prothrombin Thrombin

Fibrinogen Fibrin
5 µm
Stem Cells and the Replacement of
Cellular Elements
• The cellular elements of blood wear out and are
replaced constantly throughout a person’s life.
• Erythrocytes, leukocytes, and platelets all develop
from a common source of stem cells in the red
marrow of bones.
• The hormone erythropoietin (EPO) stimulates
erythrocyte production when oxygen delivery is low.
Differentiation of Blood Cells

Stem cells
in bone marrow

Lymphoid Myeloid
stem cells stem cells

Lymphocytes
B cells T cells

Erythrocytes Neutrophils

Platelets

Eosinophils
Monocytes Basophils
Cardiovascular Disease = Disorders of the Heart and the
Blood Vessels
• One type of cardiovascular disease, atherosclerosis, is
caused by the buildup of plaque deposits within arteries.
• A heart attack is the death of cardiac muscle tissue
resulting from blockage of one or more coronary arteries.
• A stroke is the death of nervous tissue in the brain, usually
resulting from rupture or blockage of arteries in the
brain /head.
Atherosclerosis

Connective Smooth
tissue muscle Endothelium Plaque

(a) Normal artery 50 µm (b) Partly clogged artery 250 µm

Treatment and Diagnosis of
Cardiovascular Disease
• Cholesterol is a major contributor to atherosclerosis.
• Low-density lipoproteins (LDLs) = “bad cholesterol,” are
associated with plaque formation.
• High-density lipoproteins (HDLs) = “good cholesterol,”
reduce the deposition of cholesterol.
• Hypertension = high blood pressure, promotes
atherosclerosis and increases the risk of heart attack and
stroke.
• Hypertension can be reduced by dietary changes, exercise,
and/or medication.
Adaptations for gas exchange include
pigments that bind and transport gases
• The metabolic demands of many organisms require that the
blood transport large quantities of O2 and CO2
• Blood arriving in the lungs has a low partial pressure of O2
and a high partial pressure of CO2 relative to air in the
alveoli.
• In the alveoli, O2 diffuses into the blood and CO2 diffuses
into the air.
• In tissue capillaries, partial pressure gradients favor
diffusion of O2 into the interstitial fluids and CO2 into the
blood.
Loading and unloading of respiratory gases

Alveolus Alveolus
PO2 = 100 mm Hg PCO2 = 40 mm Hg

PO2 = 40 PO2 = 100 PCO2 = 46 PCO2 = 40

Circulatory Circulatory
system system

PO2 = 40 PO2 = 100 PCO2 = 46 PCO2 = 40

PO2 ≤ 40 mm Hg PCO2 ≥ 46 mm Hg
Body tissue Body tissue
(a) Oxygen (b) Carbon dioxide
Hemoglobin
• A single hemoglobin molecule can carry four
molecules of O2
• The hemoglobin dissociation curve shows that a
small change in the partial pressure of oxygen can
result in a large change in delivery of O2
• CO2 produced during cellular respiration lowers
blood pH and decreases the affinity of hemoglobin
for O2
• This is called the Bohr shift.
  Chains

Iron
Heme

 Chains
Hemoglobin
Dissociation curves for hemoglobin at 37ºC
100
O2 unloaded

O2 saturation of hemoglobin (%)

to tissues
80
at rest

O2 unloaded
60
to tissues
during exercise
40

20

0
0 20 40 60 80 100

Tissues during Tissues Lungs

exercise at rest
PO2 (mm Hg)
(a) PO and hemoglobin dissociation at pH 7.4
2

100

O2 saturation of hemoglobin (%)

pH 7.4
80 pH 7.2

60 Hemoglobin
retains less
40 O2 at lower pH
(higher CO2
20 concentration)

0
0 20 40 60 80 100

PO2 (mm Hg)

(b) pH and hemoglobin dissociation
Membrane Structure and
Function
Overview: Life at the Edge

• The plasma membrane is the boundary that separates

the living cell from its surroundings
• The plasma membrane exhibits selective permeability,
allowing some substances to cross it more easily than
others

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Fig. 7-1
Concept 7.1: Cellular membranes are
fluid mosaics of lipids and proteins
• Phospholipids are the most abundant lipid in the plasma
membrane
• Phospholipids are amphipathic molecules, containing
hydrophobic and hydrophilic regions
• The fluid mosaic model states that a membrane is a
fluid structure with a “mosaic” of various proteins
embedded in it

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Membrane Models: Scientific Inquiry

• Membranes have been chemically analyzed and found

to be made of proteins and lipids
• Scientists studying the plasma membrane reasoned that
it must be a phospholipid bilayer

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Fig. 7-2

WATER
Hydrophilic
head

Hydrophobic
tail

WATER
• In 1935, Hugh Davson and James Danielli proposed a
sandwich model in which the phospholipid bilayer lies
between two layers of globular proteins
• Later studies found problems with this model,
particularly the placement of membrane proteins,
which have hydrophilic and hydrophobic regions
• In 1972, J. Singer and G. Nicolson proposed that the
membrane is a mosaic of proteins dispersed within the
bilayer, with only the hydrophilic regions exposed to
water

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Fig. 7-3

Phospholipid
bilayer

Hydrophobic regions Hydrophilic

of protein regions of protein
• Freeze-fracture studies of the plasma membrane
supported the fluid mosaic model
• Freeze-fracture is a specialized preparation technique
that splits a membrane along the middle of the
phospholipid bilayer

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Fig. 7-4

TECHNIQUE RESULTS
Extracellular
layer

Proteins Inside of extracellular layer

Knife

Plasma membrane Cytoplasmic layer

Inside of cytoplasmic layer
The Fluidity of Membranes

• Phospholipids in the plasma membrane can move

within the bilayer
• Most of the lipids, and some proteins, drift laterally
• Rarely does a molecule flip-flop transversely across the
membrane

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Fig. 7-5

Lateral movement Flip-flop

(~107 times per second) (~ once per month)

(a) Movement of phospholipids

Fluid Viscous

Unsaturated hydrocarbon Saturated hydro-

tails with kinks carbon tails
(b) Membrane fluidity

Cholesterol

(c) Cholesterol within the animal cell membrane

Fig. 7-5a

Lateral movement Flip-flop

(107 times per ( once per
second)of
(a) Movement month)
phospholipids
Fig. 7-6

RESULTS

Membrane proteins

Mixed proteins
after 1 hour
Mouse cell
Human cell
Hybrid cell
• As temperatures cool, membranes switch from a fluid
state to a solid state
• The temperature at which a membrane solidifies
depends on the types of lipids
• Membranes rich in unsaturated fatty acids are more
fluid that those rich in saturated fatty acids
• Membranes must be fluid to work properly; they are
usually about as fluid as salad oil

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Fig. 7-5b

Flui Viscou
d s

Unsaturated Saturated
hydrocarbon hydro-
tails
(b) with kinks
Membrane carbon tails
fluidity
• The steroid cholesterol has different effects on
membrane fluidity at different temperatures
• At warm temperatures (such as 37°C), cholesterol
restrains movement of phospholipids
• At cool temperatures, it maintains fluidity by preventing
tight packing

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Fig. 7-5c

Cholester
ol
(c) Cholesterol within the animal cell
membrane
Membrane Proteins and Their Functions

• A membrane is a collage of different proteins

embedded in the fluid matrix of the lipid bilayer
• Proteins determine most of the membrane’s specific
functions

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Fig. 7-7

Fibers of
extracellular
matrix (ECM)

Glyco- Carbohydrate
protein
Glycolipid
EXTRACELLULAR
SIDE OF
MEMBRANE

Cholesterol

Microfilaments Peripheral
of cytoskeleton proteins
Integral
protein
CYTOPLASMIC SIDE
OF MEMBRANE
• Peripheral proteins are bound to the surface of the
membrane
• Integral proteins penetrate the hydrophobic core
• Integral proteins that span the membrane are called
transmembrane proteins
• The hydrophobic regions of an integral protein consist
of one or more stretches of nonpolar amino acids, often
coiled into alpha helices

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Fig. 7-8

N-terminus EXTRACELLULAR
SIDE

C-terminus
CYTOPLASMIC
 Helix SIDE
• Six major functions of membrane proteins:
• Transport
• Enzymatic activity
• Signal transduction
• Cell-cell recognition
• Intercellular joining
• Attachment to the cytoskeleton and extracellular matrix
(ECM)

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Fig. 7-9
Signaling molecule

Enzymes Receptor

ATP
Signal transduction
(a) Transport (b) Enzymatic activity (c) Signal transduction

Glyco-
protein

(d) Cell-cell recognition (e) Intercellular joining (f) Attachment to

the cytoskeleton
and extracellular
matrix (ECM)
Fig. 7-9ac

Signaling molecule

Enzymes Receptor

ATP
Signal transduction
(a) Transport (b) Enzymatic activity (c) Signal transduction
Fig. 7-9df

Glyco-
protein

(d) Cell-cell recognition (e) Intercellular joining (f) Attachment to

the cytoskeleton
and extracellular
matrix (ECM)
The Role of Membrane Carbohydrates in Cell-Cell
Recognition

• Cells recognize each other by binding to surface

molecules, often carbohydrates, on the plasma
membrane
• Membrane carbohydrates may be covalently bonded to
lipids (forming glycolipids) or more commonly to
proteins (forming glycoproteins)
• Carbohydrates on the external side of the plasma
membrane vary among species, individuals, and even
cell types in an individual

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Synthesis and Sidedness of Membranes

• Membranes have distinct inside and outside faces

• The asymmetrical distribution of proteins, lipids, and
associated carbohydrates in the plasma membrane is
determined when the membrane is built by the ER and
Golgi apparatus

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Fig. 7-10
ER
1

Transmembrane
glycoproteins

Secretory
protein

Glycolipid

Golgi 2
apparatus

Vesicle

3
Plasma membrane:
Cytoplasmic face
4
Extracellular face
Transmembrane
Secreted glycoprotein
protein

Membrane glycolipid
Concept 7.2: Membrane structure
results in selective permeability
• A cell must exchange materials with its surroundings, a
process controlled by the plasma membrane
• Plasma membranes are selectively permeable,
regulating the cell’s molecular traffic

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The Permeability of the Lipid Bilayer

• Hydrophobic (nonpolar) molecules, such as

hydrocarbons, can dissolve in the lipid bilayer and pass
through the membrane rapidly
• Polar molecules, such as sugars, do not cross the
membrane easily

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Transport Proteins

• Transport proteins allow passage of hydrophilic

substances across the membrane
• Some transport proteins, called channel proteins, have
a hydrophilic channel that certain molecules or ions can
use as a tunnel
• Channel proteins called aquaporins facilitate the
passage of water

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• Other transport proteins, called carrier proteins, bind to
molecules and change shape to shuttle them across the
membrane
• A transport protein is specific for the substance it
moves

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Concept 7.3: Passive transport is
diffusion of a substance across a
membrane with no energy investment
• Diffusion is the tendency for molecules to spread out
evenly into the available space
• Although each molecule moves randomly, diffusion of a
population of molecules may exhibit a net movement in
one direction
• At dynamic equilibrium, as many molecules cross one
way as cross in the other direction

Animation: Membrane Selectivity Animation: Diffusion

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Fig. 7-11
Molecules of dye Membrane (cross section)

WATER

Net diffusion Net diffusion Equilibrium

(a) Diffusion of one solute

Net diffusion Net diffusion Equilibrium

Net diffusion Net diffusion Equilibrium

(b) Diffusion of two solutes

Fig. 7-11a

Molecules of dye Membrane (cross section)

WATER

Net diffusion Net diffusion Equilibrium

(a) Diffusion of one solute

• Substances diffuse down their concentration gradient,
the difference in concentration of a substance from one
area to another
• No work must be done to move substances down the
concentration gradient
• The diffusion of a substance across a biological
membrane is passive transport because it requires no
energy from the cell to make it happen

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Fig. 7-11b

Net diffusion Net diffusion Equilibrium

Net diffusion Net diffusion Equilibrium

(b) Diffusion of two solutes

Effects of Osmosis on Water Balance

• Osmosis is the diffusion of water across a selectively

permeable membrane
• Water diffuses across a membrane from the region of
lower solute concentration to the region of higher
solute concentration

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Fig. 7-12
Lower Higher Same concentration
concentration concentration of sugar
of solute (sugar) of sugar

H2O

Selectively
permeable
membrane

Osmosis
Water Balance of Cells Without Walls

• Tonicity is the ability of a solution to cause a cell to gain

or lose water
• Isotonic solution: Solute concentration is the same as
that inside the cell; no net water movement across the
plasma membrane
• Hypertonic solution: Solute concentration is greater
than that inside the cell; cell loses water
• Hypotonic solution: Solute concentration is less than
that inside the cell; cell gains water

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Fig. 7-13

Hypotonic solution Isotonic solution Hypertonic solution

H2O H2O H2O H2O

(a) Animal
cell

Lysed Normal Shriveled

H2O H2O H2O H2O

(b) Plant
cell

Turgid (normal) Flaccid Plasmolyzed

• Hypertonic or hypotonic environments create osmotic
problems for organisms
• Osmoregulation, the control of water balance, is a
necessary adaptation for life in such environments
• The protist Paramecium, which is hypertonic to its pond
water environment, has a contractile vacuole that acts
as a pump

Video: Chlamydomonas Video: Paramecium Vacuole

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Fig. 7-14
50 µm
Filling vacuole

(a) A contractile vacuole fills with fluid that enters from

a system of canals radiating throughout the cytoplasm.
Contracting vacuole

(b) When full, the vacuole and canals contract, expelling

fluid from the cell.
Water Balance of Cells with Walls

• Cell walls help maintain water balance

• A plant cell in a hypotonic solution swells until the wall
opposes uptake; the cell is now turgid (firm)
• If a plant cell and its surroundings are isotonic, there is
no net movement of water into the cell; the cell
becomes flaccid (limp), and the plant may wilt

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• In a hypertonic environment, plant cells lose water;
eventually, the membrane pulls away from the wall, a
usually lethal effect called plasmolysis

Video: Plasmolysis

Video: Turgid Elodea

Animation: Osmosis
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Facilitated Diffusion: Passive Transport
Aided by Proteins
• In facilitated diffusion, transport proteins speed the
passive movement of molecules across the plasma
membrane
• Channel proteins provide corridors that allow a specific
molecule or ion to cross the membrane
• Channel proteins include
• Aquaporins, for facilitated diffusion of water
• Ion channels that open or close in response to a stimulus
(gated channels)

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Fig. 7-15
EXTRACELLULAR
FLUID

Channel protein Solute

CYTOPLASM

(a) A channel protein

Carrier protein Solute

(b) A carrier protein

• Carrier proteins undergo a subtle change in shape that
translocates the solute-binding site across the
membrane

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• Some diseases are caused by malfunctions in specific
transport systems, for example the kidney disease
cystinuria

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Concept 7.4: Active transport uses
energy to move solutes against their
gradients
• Facilitated diffusion is still passive because the solute
moves down its concentration gradient
• Some transport proteins, however, can move solutes
against their concentration gradients

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The Need for Energy in Active
Transport
• Active transport moves substances against their
concentration gradient
• Active transport requires energy, usually in the form of
ATP
• Active transport is performed by specific proteins
embedded in the membranes

Animation: Active Transport

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• Active transport allows cells to maintain concentration
gradients that differ from their surroundings
• The sodium-potassium pump is one type of active
transport system

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Fig. 7-16-1

EXTRACELLULAR [Na+] high

FLUID [K+] low

Na+

Na+

[Na+] low
Na+
CYTOPLASM [K+] high

1 Cytoplasmic Na+ binds

to
the sodium-potassium
pump.
Fig. 7-16-2

Na+

Na+

Na+

ATP
P
ADP

2 Na+ binding stimulates

phosphorylation by ATP.
Fig. 7-16-3

Na+
Na+

Na+

3 Phosphorylation causes
the protein to change its
shape. Na+ is expelled to
the outside.
Fig. 7-16-4

K+

K+

P
P
4 K+ binds on the
extracellular side and
triggers release of the
phosphate group.
Fig. 7-16-5

K+

K+

5 Loss of the phosphate

restores the protein’s original
shape.
Fig. 7-16-6

K+

K+

6 K+ is released, and the

cycle repeats.
Fig. 7-16-7

EXTRACELLULAR
[Na+] high Na+
FLUID [K+] low Na+

Na+ Na+ Na+

Na+ Na+

Na+

[Na+] low ATP

Na+ P
[K+] high P
CYTOPLASM ADP
1 2 3

K+

K+

+
K+
K
K+

P
K+ P
6 5 4
Fig. 7-17
Passive transport Active transport

ATP
Diffusion Facilitated diffusion
How Ion Pumps Maintain Membrane
Potential
• Membrane potential is the voltage difference across a
membrane
• Voltage is created by differences in the distribution of
positive and negative ions

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• Two combined forces, collectively called the
electrochemical gradient, drive the diffusion of ions
across a membrane:
• A chemical force (the ion’s concentration gradient)
• An electrical force (the effect of the membrane potential on
the ion’s movement)

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• An electrogenic pump is a transport protein that
generates voltage across a membrane
• The sodium-potassium pump is the major electrogenic
pump of animal cells
• The main electrogenic pump of plants, fungi, and
bacteria is a proton pump

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Fig. 7-18

– EXTRACELLULAR
+
FLUID
ATP – + H+

H+
Proton pump
H+

– + H+
H+
– +
CYTOPLASM
H+
– +
Cotransport: Coupled Transport by a
Membrane Protein
• Cotransport occurs when active transport of a solute
indirectly drives transport of another solute
• Plants commonly use the gradient of hydrogen ions
generated by proton pumps to drive active transport of
nutrients into the cell

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Fig. 7-19

– +
ATP H+
H+
– +
Proton pump H+
H+

– +
H+ – H+
+
H+ Diffusion
of H+
Sucrose-H+
cotransporter
H+

Sucrose – +

– + Sucrose
Concept 7.5: Bulk transport across the
plasma membrane occurs by exocytosis
and endocytosis
• Small molecules and water enter or leave the cell
through the lipid bilayer or by transport proteins
• Large molecules, such as polysaccharides and proteins,
cross the membrane in bulk via vesicles
• Bulk transport requires energy

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Exocytosis

• In exocytosis, transport vesicles migrate to the

membrane, fuse with it, and release their contents
• Many secretory cells use exocytosis to export their
products

Animation: Exocytosis
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Endocytosis

• In endocytosis, the cell takes in macromolecules by

forming vesicles from the plasma membrane
• Endocytosis is a reversal of exocytosis, involving
different proteins
• There are three types of endocytosis:
• Phagocytosis (“cellular eating”)
• Pinocytosis (“cellular drinking”)
• Receptor-mediated endocytosis

Animation: Exocytosis and Endocytosis Introduction

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• In phagocytosis a cell engulfs a particle in a vacuole
• The vacuole fuses with a lysosome to digest the particle

Animation: Phagocytosis
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Fig. 7-20
PHAGOCYTOSIS
EXTRACELLULAR CYTOPLASM 1 µm
FLUID
Pseudopodium
Pseudopodium
of amoeba

“Food”or
other particle Bacterium
Food
vacuole Food vacuole
An amoeba engulfing a bacterium
via phagocytosis (TEM)

PINOCYTOSIS

0.5 µm
Plasma
membrane Pinocytosis vesicles
forming (arrows) in
a cell lining a small
blood vessel (TEM)

Vesicle

RECEPTOR-MEDIATED ENDOCYTOSIS
Coat protein
Receptor Coated
vesicle

Coated
pit
Ligand

A coated pit
Coat and a coated
protein vesicle formed
during
receptor-
mediated
endocytosis
(TEMs)

Plasma
membrane
0.25 µm
Fig. 7-20a

PHAGOCYTOSIS
EXTRACELLULAR CYTOPLASM 1 µm
FLUID
Pseudopodium
Pseudopodium
of amoeba

“Food” or
other particle Bacterium
Food
vacuole Food vacuole
An amoeba engulfing a bacterium
via phagocytosis (TEM)
• In pinocytosis, molecules are taken up when
extracellular fluid is “gulped” into tiny vesicles

Animation: Pinocytosis

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Fig. 7-20b

PINOCYTOSIS

0.5 µm
Plasma
membrane Pinocytosis vesicles
forming (arrows) in
a cell lining a small
blood vessel (TEM)

Vesicle
• In receptor-mediated endocytosis, binding of ligands to
receptors triggers vesicle formation
• A ligand is any molecule that binds specifically to a
receptor site of another molecule

Animation: Receptor-Mediated Endocytosis

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Fig. 7-20c
RECEPTOR-MEDIATED ENDOCYTOSIS

Coat protein
Receptor Coated
vesicle

Coated
pit
Ligand

A coated pit
Coat and a coated
protein vesicle formed
during
receptor-
mediated
endocytosis
(TEMs)

Plasma
membrane
0.25 µm
Fig. 7-UN1
Passive transport:
Facilitated diffusion

Channel Carrier
protein protein
Fig. 7-UN2
Active transport:

ATP
Fig. 7-UN3

Environment:
“Cell” 0.01 M sucrose
0.03 M sucrose 0.01 M glucose
0.02 M glucose 0.01 M fructose
Fig. 7-UN4
You should now be able to:

1. Define the following terms: amphipathic molecules,

aquaporins, diffusion
2. Explain how membrane fluidity is influenced by
temperature and membrane composition
3. Distinguish between the following pairs or sets of
terms: peripheral and integral membrane proteins;
channel and carrier proteins; osmosis, facilitated
diffusion, and active transport; hypertonic, hypotonic,
and isotonic solutions

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4. Explain how transport proteins facilitate diffusion
5. Explain how an electrogenic pump creates voltage
across a membrane, and name two electrogenic
pumps
6. Explain how large molecules are transported across a
cell membrane

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin

Cummings
Bioelectricity
Membrane Potentials
• A. The body as a whole is electrically neutral
• B. All of the cells of body have an electrical potential across
their membrane (Voltage difference) known as the
membrane potential
• C. Membrane potentials develop because of differing ion
concentrations between the inside and outside of the cell
Membrane Potentials
Membrane Potentials
• Principals of electricity - Potential difference is determined
by the difference in charge between two points
• 1. Units of electrical potential are in volts (V) or for
biological system millivolts (mV) 1 V = 1000mV
• 2. Voltage is always measured between two points
(Potential difference)
Membrane Potentials
• B. Current - flow of electrical charges from one point to
another
• 1. Like charges repel unlike attract
• 2. Ions tend to move from areas of greater concentration to
areas of least concentration
• 3. Movement of a positive ion from one side of a
membrane to the other implies a negative charge is left
behind
Membrane Potentials
• C. Current Flow
• Ohm’s Law - I = E / R, R = resistance
• I = current flow, E = electrical potential
• 1. Cell - Aqueous solution + good conductor (Ions and water)
• 2. Lipid membrane - A few charged groups can not carry current - high
electrical resistance - good insulator
• 3. ECF and ICF - both have low electrical resistance
Membrane Potentials
• Resting Membrane Potential
• 1. By convention - ECF (outside of the cell) is
assigned a voltage of zero
• 2. Polarity of the membrane is stated in terms of
the sign of the excess charge inside of the cell
 Membrane Ion Channels
• Types of Channels
• 1. Leak channels - Open all of the time - slow leak of
ions
• a. Sodium, potassium & chlorine
• b. Membrane 75% more permeable to K+ than Na+
• c. Accounts for 95% of the resting membrane
potential
Membrane Ion Channels
• 2. Na+K+ATPase Pump
• a. Unequal transport of positive ions makes the ICF more negative
than it would be from diffusion alone - 2 K+ inside and 3 Na+ to
outside
• b. Electrogenic pump
• c. Accounts for 5% of resting membrane potential
Membrane Potentials
Ion Gradients
• The ion gradients have two forms.
• 1. Chemical Concentration Gradient
• 2. Electrical concentration gradient
• (Charge buildup and charge differential)
• Together these form what is known as the
electrochemical gradient
Resting Membrane Potential
• 1. In all cells a potential difference across the membrane exists
• a. Inside is negative (Na+K+ATPase)
• b. Membrane potentials usually within -40 to -90 mv
• 2. A cell with a resting membrane potential is said to be polarized
• 3. Both the inside and the outside of the cell are electrically neutral
Resting Membrane Potential
• B. Factors that determine the resting membrane potential
• Selective permeability of the of the plasma
membrane
• Leak channels
• Na+K+ATPase pump
• Differences in ion concentrations
Membrane Potentials

IONS INSIDE OUTSIDE

Na+ 14 140

K+ 140 4

Cl- 4 108
Resting Membrane Potential
• 2. Many substances are in the cell but the mobile ions Na+, K+, Ca++
and Cl- play the most important roles
• 3. ECF - Cl- helps to balance Na+
• ICF - Proteins (Neg charge) balance K+
Resting Membrane Potential
• 4. Selective membrane permeability
• a. At rest - Slightly permeable to Na+, 75 times
more permeable to K+, and freely permeable to
Cl-
• b. K+ moves down it’s concentration gradient
more easily & faster than Na+
• c. Movement of a K+ out leaves a negative
charge behind
Resting Membrane Potential
• d. Why no equilibrium?
• Na+K+ATPase pump - stabilizes resting
membrane potential by maintaining diffusion
gradients for Na+ and K+
• Concentration gradient – Limit to ability of
Na+K+ATPase pump
• c. Cl- Movement out = movement in - no contribution to
membrane potential
Equilibrium Potential

• Equilibrium potential or electrochemical

potential at which ion movements in both
directions across the membrane are exactly
balanced (net movement = zero)
• 1. Ion flux = 0 implies no net ion movement
• 2. The value of the equilibrium potential (Nernst potential)
for any ion depends on the concentration gradient across
the membrane for that ion
Equilibrium Potential
• 4. The greater the concentration gradient the greater the
equilibrium potential
• 5. The equilibrium potential for one ion can be different in
magnitude and direction from those of other ions
• 6. Given the ion concentration gradient the Nernst
potential for any ion can be calculated. The Nernst
equation is used to determine the electrochemical potential
for any ion across the biological membrane.
Equilibrium Potential
• Nernst Equation

• E(x) = RT/ZF log [x]inside/[x]outside

• R = Gas constant
• T = Temp. degrees Kelvin
• Z = Charge on ion (Valance)
• F = Faraday’s constant
Membrane Potentials
Nernst’s Equation
• A more useful form of the Nernst’s equation is -

• E(x) = -61mV log [X]inside / [X]outside

• or
• = 61 mV log [X]outside / [X]inside
Membrane Potentials
Nernst’s Equation - Examples
• Example 1 - Calculate the electrochemical potential for Na+

• E Na+ = -61mV log [14]/[140]

• log of 0.10 = -1
• then
• E Na+ = 61 mV
Membrane Potentials
Nernst’s Equation - Examples
• Example 2 - Calculate the electrochemical potential for K+

• E K+ = -61mV log [140]/[4]

• log of 35 = 1.5441
• then
• E K+ = - 94 mV
Membrane Potentials
Nernst Equation - Examples
Membrane Potentials
Resting Membrane Potential
• In reality a living cell contains a great number of ionic species. Most
of these can and do move in and out of the cell others such as
proteins can not without help. The net movement of all ionic
currents across the membrane determines the resting membrane
potential.
Membrane Potentials
Resting Membrane Potential
• The net current flow ( I ) across the membrane is given by;

• I(x) = g(x) {Em - E(x)}

• Where - g(x) is ion conductance

• Em is resting membrane Pot.
• E(x) is the Nernst’s Pot.
Membrane Potentials
Resting Membrane Potential
• At rest the membrane potential is not changing, then the sum of all
currents must equal zero.
• Thus
• I Na+ + I K+ + I Cl- + … = 0
Membrane Potentials
Resting Membrane Potential
• Therefore
• g Na+[Em -ENa+] + g K+[Em - EK+] + g Cl-[Em - ECl-] + … = 0
Membrane Potentials
Resting Membrane Potential
• Solving for Em yields the Goldman equation which gives the resting
membrane potential
Membrane Potentials
Resting Membrane Potential
• Goldman Equation
• Em = [gNa+/(gNa+ + gK+ + gCl- )] E Na+
• + [gK+/(gNa+ + gK+ + gCl- )] E K+
• + [gCl-/(gNa+ + gK+ + gCl- )] E Cl-
• +…
• Thus the resting membrane potential is a summation of all of the ion
potentials times their percentage of the total ion conductance
Membrane Potentials
Resting Membrane Potential
• Since K+ conductance is almost 75 times that of Na+. The
resting membrane potential is much closer to the Nernst’s
potential for K+ than it is to the Nernst’s potential for Na+.
• Why would K+ conductance predominate?
Excitable Cells
• A. Nerve and muscle cells are excitable (Em < -40 mV)
• 1. Electrochemical impulses are transient and
rapid changes in Em
• 2. Two forms of electrochemical impulses
• B. Electrochemical signals
• 1. Graded potentials - short distance
• 2. Action potentials - long distance
A typical neuron

(Note – the term ‘nerve’ can refer to collections of neurons)

The giant axon of squid
• Axons up to 1mm in diameter – 1000 times that of mammalian nerves

• Hodgkin and Huxley (1939) measured resting potential

• Hodgkin and Katz (1949) measured change in potential resulting from

manipulating K+ concentration
• Permeability to potassium is the primary source of the resting membrane
potential (NB permeability due to different process from gates)
Ionic gradients maintain electrical potentials
• Electrical potentials across cell membranes are generated by
differences in concentration of specific ions (K+, Na+, Cl-) which are
maintained by ion transporters (use energy for energetically
unfavourable reaction)
Absolute temp Permeability of membrane to ion

Gas constant
RT   Pi [ Ai ]2   Pi [ Ai ]1 
  Concentration of ion
V ln  
Voltage F   Pi [ Ai ]1   Pi [ Ai ]2 

Faraday constant
Ratio of outside to inside
Goldman equation (from Nernst equation)
Ion Intracellular Extracellular
Potassium 400 20
Concentrations in a squid giant axon –
Sodium 50 440
the resting potential is -65mV
Chloride 4-150 560
Calcium 0.0001 10
Electrical signals are generated by ionic
movement

• The resting potential is maintained by active transport of ions (e.g. the Na/K pump – 2 K+
pumped in and 3 Na+ out for one ATP).

• Electrical signals occur when selective ion channels open to allow specific ions to move down
gradient

• Depolarisation largely caused by change in permeability to Na + ions

Action potentials in the squid giant axon
 Electrical signals are propagated by voltage-dependent channels

• A threshold depolarisation causes adjacent sodium channels to open

• Signals move along axons

The properties of single ion channels can be observed and manipulated using the patch-clamp method
Myelinated nerve cells have much faster rates of transmission

Passive diffusion of Na+ ions triggers voltage-dependent gates at the next node of Ranvier

Increases speed of action potential from 1m/s to about 18 m/s in mammalian nerves
Nerve cells communicate via synapses
• Electrical synapses via gap junctions between cells allow direct passage of
electrical signal

• Chemical synapses enable communication by the release of

neurotransmitters
• Neurotransmitters in pre-synaptic vesicles
• Release triggered by Ca2+ influx through voltage-gated channels
• Chemical diffuse across synapse and recognised by receptors
• Threshold response in post-synaptic nerve
• Computation through pre- and post-synaptic modification

• A similar process allows communication between nerves and effectors

• e.g. acetylcholine released at nerve-muscle end plates
The diversity of neurotransmitters
• Over 100 neurotransmitters

• Two classes of neurotransmitter

• Neuropeptides: 3-36 amino acid peptides such as - and -endorphins
• Small-molecule neurotransmitters including amino acids (glutamate, aspartate), purines (ATP)
and biogenic amines such as serotonin, dopamine and histamine

• Examples
• Acetylcholine at neuromuscular junctions
• Glutamate is the most important transmitter for normal brain function

• Drugs and disease

• Most psychotrophic drugs alter steps in the generation and release of neurotransmitters (e.g.
fluoxetine – Prozac – blocks reuptake of serotonin)
 Nerve cells are stimulated by diverse receptor mechanisms

• Somatic sensory receptors

• E.g. Muscle spindles, Merkel’s discs, Meissner’s corpuscles
• Pain receptors
• Visual/photoperiod receptors
• Retina, pineal gland
• Auditory receptors
• Cochlea
• The vestibular system
• Otolith organs (utricle and sacculus) and semicircular canals of the inner ear
• Chemical receptors
• Olfaction, taste, trigeminal chemsonsory system
• The conversion of signal to nervous response is called signal transduction
Hodgkin-Huxley Model
and
FitzHugh-Nagumo Model
Nervous System
• Signals are propagated from nerve cell to nerve cell
(neuron) via electro-chemical mechanisms
• ~100 billion neurons in a person
• Hodgkin and Huxley experimented on squids and
discovered how the signal is produced within the
neuron
• H.-H. model was published in Jour. of Physiology
(1952)
• H.-H. were awarded 1963 Nobel Prize
• FitzHugh-Nagumo model is a simplification
Neuron

C. George Boeree: [Link]/~cgboeree/

Action Potential
mV
Axon membrane
_ 30 potential difference
V = Vi – Ve
When the axon is
_0
excited, V spikes
V because sodium
Na+ and potassium
K+ ions flow
through the
membrane.
10 msec
-70
 Nernst Potential
VNa , VK and Vr

Ion flow due to

electrical signal

Traveling wave

C. George Boeree: [Link]/~cgboeree/

 Circuit Model for Axon Membrane
Since the membrane separates charge, it is modeled as a
capacitor with capacitance C. Ion channels are resistors.
1/R = g = conductance

iC = C dV/dt

iNa = gNa (V – VNa)

iK= gK (V – VK)

ir = gr (V – Vr)
 Circuit Equations
Since the sum of the currents is 0, it follows that

dV
C   g Na (V  V Na )  g K (V  V K )  gr(V  Vr )  Iap
dt
where Iap is applied current. If ion conductances are constants
then group constants to obtain 1st order, linear eq

dV
C   g (V  V *)  Iap
dt
Solving gives
V (t )  V *  Iap / g
Variable Conductance
g

Experiments showed that gNa and gK varied with time and V. After
stimulus, Na responds much more rapidly than K .
 Hodgkin-Huxley System
Four state variables are used:
v(t)=V(t)-Veq is membrane potential,
m(t) is Na activation,
n(t) is K activation and
h(t) is Na inactivation.

In terms of these variables gK=gKn4 and gNa=gNam3h.

The resting potential Veq≈-70mV.
≈-70mV Voltage clamp experiments
determined gK and n as functions of t and hence the parameter
dependences on v in the differential eq. for n(t). Likewise for m(t)
and h(t).
Hodgkin-Huxley System
dv
C   g Na m h(v  VNa )  g K n (v  VK )  gr (v  Vr )  I ap
3 4

dt

dm
 m( v )(1  m )   m( v )m
dt
dn
 n ( v )(1  n )   n ( v )n
dt
dh
 h ( v )(1  h )   h ( v )h
dt
110 mV
Iap =8, v(t)

1.2
m(t)

n(t)

40msec
h(t)

10msec

Iap=7, v(t)
 Fast-Slow Dynamics
m(t)
ρm(v) dm/dt = m∞(v) – m.
ρm(v) is much smaller than
n(t) ρn(v) and ρh(v). An increase in
v results in an increase in
m∞(v) and a large dm/dt.
h(t) Hence Na activates more
rapidly than K in response to a
change in v.
10msec

v, m are on a fast time scale and n, h are slow.

 FitzHugh-Nagumo System

dv dw
  f (v )  w  I and  v  0.5w
dt dt
I represents applied current, ε is small and f(v) is a cubic nonlinearity. Observe that in the (v,w)
phase plane
dw  ( v  0.5 w )

dv f (v )  w  I
which is small unless the solution is near f(v)-w+I=0. =0 Thus the slow manifold is the cubic w=f(v)
+I which is the nullcline of the fast variable v. And w is the slow variable with nullcline w=2v.
 Take f(v)=v(1-v)(v-a) .

Stable rest state I=0 Stable oscillation I=0.2

w w

v v
Web resources
• [Link]
 Molecular motors

Molecular motors are a class of proteins that drive intracellular trafficking by converting
chemical energy to mechanical work along cytoskeletal filaments.
Examples:
• Flagellar Motors
• Actin –Myosin Complexes
• Proton Pumps
Actin-based Motor Proteins Are Members of the Myosin Superfamily
Perhaps the most fascinating proteins that associate with the cytoskeleton are the molecular
motors called motor proteins. These remarkable proteins bind to a polarized cytoskeletal
filament and use the energy derived from repeated cycles of ATP hydrolysis to move steadily
along it. Dozens of different motor proteins coexist in every eucaryotic cell. They differ in
the type of filament they bind to (either actin or microtubules), the direction in which they
move along the filament, and the “cargo” they carry. Many motor proteins carry membrane-
enclosed organelles—such as mitochondria, Golgi stacks, or secretory vesicles—to their
appropriate locations in the cell. Other motor proteins cause cytoskeletal filaments to slide
against each other, generating the force that drives such phenomena as muscle contraction,
ciliary beating, and cell division.

Actin
Abundant protein that forms actin filaments in all eucaryotic cells. The monomeric form is
sometimes called globular or G-actin; the polymeric form is filamentous or F-actin.
The first motor protein identified was skeletal muscle myosin, which is
responsible for generating the force for muscle contraction. This myosin,
called myosin II (see below) is an elongated protein that is formed from two heavy
chains and two copies of each of two light chains. Each of the heavy chains has a
globular head domain at its N-terminus that contains the force-generating
machinery, followed by a very long amino acid sequence that forms an
extended coiled-coil that mediates heavy chain dimerization
Some myosins (such as VIII and XI) have been found only in plants, and some have been
found only in vertebrates (IX). Most, however, are found in all eucaryotes, suggesting that
myosins arose early in eucaryotic evolution. The yeast Saccharomyces cerevisiae contains
five myosins: two myosin Is, one myosin II, and two myosin Vs. One can speculate that
these three types of myosins are necessary for a eucaryotic cell to survive and that other
myosins perform more specialized functions in multicellular organisms. The nematode C.
elegans, for example, has at least 15 myosin genes, representing at least seven structural
classes; the human genome includes about 40 myosin genes.
Muscle contraction is the most familiar and the best understood form of
movement in animals. In vertebrates, running, walking, swimming, and
flying all depend on the rapid contraction of skeletal muscle on its
scaffolding of bone, while involuntary movements such as heart pumping
and gut peristalsis depend on the contraction of cardiac muscle and
smooth muscle, respectively. All these forms of muscle contraction
depend on the ATP-driven sliding of highly organized arrays
of actin filaments against arrays of myosin II filaments.
A myosin II molecule is composed of two heavy chains (each about 2000 amino acids long (green) and four light
chains (blue). The light chains are of two distinct types, and one copy of each type is present on each myosin head.
Dimerization occurs when the two α helices of the heavy chains wrap around each other to form a coiled-coil,
driven by the association of regularly spaced hydrophobic amino acids (see Figure 3-11). The coiled-coil
arrangement makes an extended rod in solution, and this part of the molecule is called the tail.

All of the myosins except one move toward the plus end of an actin filament, although they do so at different
speeds.
Microtubules are small tubes with a diameter of 25nm, built from
smaller building blocks of tubulin. They are a key component of the
cytoskeleton where, because of their long persistence length of several
mm due to the high bending rigidity of around 30×10−24Nm2 [13], they
play an important role where mechanical stability is required in cells.
Examples are the dendritic structures in axons, mitotic spindles during
cell division and of course the axoneme.
There Are Two Types of Microtubule Motor Proteins: Kinesins and Dyneins

Motor domains of kinesins are also conserved

Kinesins is a motor protein that moves along microtubules. It was first identified in the
giant axon of the squid, where it carries membrane-enclosed organelles away from the
neuronal cell body toward the axon terminal by walking toward the plus end of
microtubules. Kinesin is similar structurally to myosin II in having two heavy chains and
two light chains per active motor, two globular head motor domains, and an
elongated coiled-coil responsible for heavy chain dimerization. Like myosin, kinesin is a
member of a large protein superfamily, for which the motor domain is the only common
element
The dyneins are a family of minus-end-directed microtubule motors, but they
are unrelated to the kinesin superfamily. They are composed of two or three
heavy chains (that include the motor domain) and a large and variable
number of associated light chains. The dynein family has two major
branches. The most ancient branch contains the cytoplasmic dyneins, which
are typically heavy-chain homodimers, with two large motor domains as
heads. Cytoplasmic dyneins are probably found in all eucaryotic cells, and
they are important for vesicle trafficking, as well as for localization of the
Golgi apparatus near the center of the cell. Axonemal dyneins, the other large
branch, include heterodimers and heterotrimers, with two or three motor-
domain heads, respectively.
Comparison of the
mechanochemical
cycles of kinesin and
myosin II
Cilia and flagella are actively bending hair-like appendages that act as sensing
and motility generating organelles of eukaryotic cells. Their evolutionary
highly conserved working mechanism as well as their widespread occurrence
in a great variety of systems demonstrate the power and importance of
physical interactions as a means of achieving biological function.

Prokaryotic flagella of e.g. E. coli

Cilia and Flagella Are Motile Structures Built from
Microtubules and Dyneins
•Wave like movement of cilia and beat like movement of
flagellum

Arrangement of microtubules in a flagellum or cilium.

Ciliary dynein. Ciliary (axonemal) dynein is a large protein assembly (nearly 2 million
daltons) composed of 9–12 polypeptide chains, the largest of which is the heavy chain of
more than 500,000 daltons.
The axoneme consists of nine cylindrically arranged
microtubule doublets, that are connected by the
molecular motor protein dynein

The bending of an axoneme

(A) When axonemes are exposed to
the proteolytic enzyme trypsin, the
linkages holding neighboring doublet
microtubules together are broken. In this
case, the addition of ATP allows the
motor action of the dynein heads to slide
one pair of doublet microtubules against
the other pair. (B) In an
intact axoneme (such as in a sperm),
sliding of the doublet microtubules is
prevented by flexible protein links. The
motor action therefore causes a bending
motion, creating waves or beating
motions,
The main electrogenic pump of plants, fungi, and bacteria is a
proton pump
ATP
synthase
•The ATP synthase enzyme
is reversible and can also
serve as a proton pump by
coupling ATP hydrolysis to
proton translocation. Each
of the respiratory enzymes
uses a different strategy for
performing proton
pumping.
Conformational couplings in protonation-coupled membrane transporters. Proton pumps use an external energy source to
transfer protons against the electrochemical ion gradient; by contrast, proton channels open in response to an external input,
such as ligand binding, to allow protons to pass down their electrochemical gradient.

The proton pump (H+/K+-ATPase) is the final common pathway for acid secretion in gastric parietal cells, and inhibition
of the pump blocks acid secretion almost completely. Proton pump inhibitors are pro-drugs that are rapidly absorbed
from the small intestine.
Proton pumps, bacteriorhodopsin and ATP synthases in particular, are capable
of continuous, renewable conversion of light to chemical, mechanical or
electrical energy, which can be used in macro- or nano-scale devices. The
capability of protein systems incorporated into liposomes to generate ATP,
which can be used to drive chemical reactions and to act as molecular motors
has been already demonstrated. Other possible applications of such
biochemical devices include targeted drug delivery and biocatalytic reactors.
All these devices might prove superior to their inorganic alternatives.
References

[Link]
%20proteins,to%20move%20steadily%20along%20it.
 FRAP FCS
Fluorescence correlation spectroscopy (FCS)
And
fluorescence recovery after photobleaching (FRAP)

are widely used methods to determine diffusion coefficients

FRAP

FRAP is a well-known fluorescence microscopy technique that has been around since the 1970s (Axelrod et al. 1976;
Peters et al. 1974). FRAP allows to measure the diffusion of fluorescently labeled molecules or particles on a micrometer
scale. A typical FRAP experiment consists of three distinct phases, First, with a low-intensity excitation beam, the
fluorescence signal is measured coming from the region of interest in the fluorescently labeled sample. Next, with a high-
power excitation beam, the fluorescent molecules are quickly photobleached in a particular area, typically in the order of
a few micrometer up to tens of micrometers in diameter.
In the drug delivery field, FRAP was used to study the mobility of macromolecules and
nanomedicines in extracellular matrices, such as mucus, (tumor) cell interstitium , and
vitreous. For example, by studying the mobility of differently sized macromolecules in lung
mucus and bovine vitreous, researchers have found that the hyaluronic acid network in the
interfibrillar spaces of vitreous poses an extra-sterical hindrance on the diffusing molecules as
a function of their size, which is not the case for lung mucus (Braeckmans et al. 2003).

FRAP has been used to study the mobility of nucleotide acids in living cells.

In related pharmaceutical research, FRAP has been used for studying gel systems for time-
controlled drug release as well
Fluorescence Correlation Spectroscopy

FCS is a powerful complementary technique to FRAP that was developed during the same period an FCS experiment is
based on a CLSM type of instrument such that only light from the focal spot can reach the detector. However, rather
than being scanned across the sample for obtaining an image, here the focused laser beam is held stationary at one
particular location of interest in the sample. The fluorescence intensity is monitored with very high sensitivity and
temporal resolution using avalanche photo diode detectors.
FCS can be used to study photodynamics of fluorophores, as well as binding equilibria and kinetics of enzymes,
proteins, and nucleic acids.

Dual-color FCS is a suitable method for studying the integrity of liposomal siRNA formulations in biological fluids, such
as full human serum.

Dual-color FCS technique to study the association and dissociation of antisense oligonucleotides and cationic polymers
in buffer and of antisense oligonucleotides and cationic liposomes in living cells