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Chapter 42

Circulation and Gas Exchange


PowerPoint Lectures for Biology, Seventh Edition
Neil Campbell and Jane Reece

MATERIALS FOR BIOLOGY OLYMPIAD PREPARATION DWIWARNA HIGH BOARDING SCHOOL, BOGOR INSTRUCTOR: WIDIATI and SUGANDA KUSMANA CORRESPONDENCE: suganda_kusmana@yahoo.com

Lectures by Chris Romero


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Overview: Trading with the Environment Every organism must exchange materials with its environment
And this exchange ultimately occurs at the cellular level

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In unicellular organisms
These exchanges occur directly with the environment

For most of the cells making up multicellular organisms


Direct exchange with the environment is not possible

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The feathery gills projecting from a salmon


Are an example of a specialized exchange system found in animals

Figure 42.1
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Concept 42.1: Circulatory systems reflect phylogeny Transport systems


Functionally connect the organs of exchange with the body cells

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Most complex animals have internal transport systems


That circulate fluid, providing a lifeline between the aqueous environment of living cells and the exchange organs, such as lungs, that exchange chemicals with the outside environment

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Invertebrate Circulation The wide range of invertebrate body size and form
Is paralleled by a great diversity in circulatory systems

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Gastrovascular Cavities Simple animals, such as cnidarians


Have a body wall only two cells thick that encloses a gastrovascular cavity

The gastrovascular cavity


Functions in both digestion and distribution of substances throughout the body

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Some cnidarians, such as jellies


Have elaborate gastrovascular cavities
Circular canal

Mouth

Radial canal 5 cm Figure 42.2

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Open and Closed Circulatory Systems More complex animals


Have one of two types of circulatory systems: open or closed

Both of these types of systems have three basic components


A circulatory fluid (blood) A set of tubes (blood vessels)

A muscular pump (the heart)

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In insects, other arthropods, and most molluscs


Blood bathes the organs directly in an open circulatory system
Heart

Hemolymph in sinuses surrounding ograns

Anterior vessel

Lateral vessels

Ostia

Figure 42.3a

Tubular heart (a) An open circulatory system

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In a closed circulatory system


Blood is confined to vessels and is distinct from the interstitial fluid
Heart

Interstitial fluid

Small branch vessels in each organ

Dorsal vessel (main heart)

Auxiliary hearts

Ventral vessels

Figure 42.3b

(b) A closed circulatory system

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Closed systems
Are more efficient at transporting circulatory fluids to tissues and cells

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Survey of Vertebrate Circulation Humans and other vertebrates have a closed circulatory system
Often called the cardiovascular system

Blood flows in a closed cardiovascular system


Consisting of blood vessels and a two- to fourchambered heart

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Arteries carry blood to capillaries


The sites of chemical exchange between the blood and interstitial fluid

Veins
Return blood from capillaries to the heart

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Fishes A fish heart has two main chambers


One ventricle and one atrium

Blood pumped from the ventricle


Travels to the gills, where it picks up O2 and disposes of CO2

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Amphibians Frogs and other amphibians


Have a three-chambered heart, with two atria and one ventricle

The ventricle pumps blood into a forked artery


That splits the ventricles output into the pulmocutaneous circuit and the systemic circuit

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Reptiles (Except Birds) Reptiles have double circulation


With a pulmonary circuit (lungs) and a systemic circuit

Turtles, snakes, and lizards


Have a three-chambered heart

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Mammals and Birds In all mammals and birds


The ventricle is completely divided into separate right and left chambers

The left side of the heart pumps and receives only oxygen-rich blood
While the right side receives and pumps only oxygen-poor blood

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A powerful four-chambered heart


Was an essential adaptation of the endothermic way of life characteristic of mammals and birds

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Vertebrate circulatory systems

FISHES

AMPHIBIANS

REPTILES (EXCEPT BIRDS)

MAMMALS AND BIRDS

Gill capillaries

Lung and skin capillaries

Lung capillaries

Lung capillaries

Artery

Gill circulation

Pulmocutaneous circuit

Heart: ventricle (V) A Atrium (A) Systemic Vein circulation V Left Right Systemic circuit A

Right systemic aorta


A V Right

Pulmonary circuit Left Systemic V aorta Left A

Pulmonary circuit

A V Right

A V Left Systemic circuit

Systemic capillaries

Systemic capillaries
Figure 42.4

Systemic capillaries

Systemic capillaries

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Concept 42.2: Double circulation in mammals depends on the anatomy and pumping cycle of the heart The structure and function of the human circulatory system
Can serve as a model for exploring mammalian circulation in general

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Mammalian Circulation: The Pathway Heart valves


Dictate a one-way flow of blood through the heart

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Blood begins its flow


With the right ventricle pumping blood to the lungs

In the lungs
The blood loads O2 and unloads CO2

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Oxygen-rich blood from the lungs


Enters the heart at the left atrium and is pumped to the body tissues by the left ventricle

Blood returns to the heart


Through the right atrium

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The mammalian cardiovascular system


7

Anterior vena cava

Capillaries of head and forelimbs

Pulmonary artery Capillaries of right lung

Aorta
9 6

Pulmonary artery

2 3 4 11 3

Capillaries of left lung

Pulmonary vein Right atrium

1 10

Left atrium

Pulmonary vein

Left ventricle Right ventricle Aorta

Posterior vena cava


8

Capillaries of abdominal organs and hind limbs

Figure 42.5
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The Mammalian Heart: A Closer Look A closer look at the mammalian heart
Provides a better understanding of how double circulation works
Pulmonary artery Aorta Pulmonary artery Left atrium Pulmonary veins Anterior vena cava Right atrium

Pulmonary veins

Semilunar valve Atrioventricular valve

Semilunar valve Atrioventricular valve

Posterior vena cava

Figure 42.6

Right ventricle Left ventricle

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The heart contracts and relaxes


In a rhythmic cycle called the cardiac cycle

The contraction, or pumping, phase of the cycle


Is called systole

The relaxation, or filling, phase of the cycle


Is called diastole

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The cardiac cycle


Semilunar valves closed
2 Atrial systole; ventricular diastole

0.1 sec

0.3 sec 0.4 sec

Semilunar valves open

AV valves open

1 Atrial and ventricular diastole

AV valves closed 3 Ventricular systole; atrial diastole

Figure 42.7
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The heart rate, also called the pulse


Is the number of beats per minute

The cardiac output


Is the volume of blood pumped into the systemic circulation per minute

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Maintaining the Hearts Rhythmic Beat Some cardiac muscle cells are self-excitable
Meaning they contract without any signal from the nervous system

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A region of the heart called the sinoatrial (SA) node, or pacemaker


Sets the rate and timing at which all 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
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The impulses that travel during the cardiac cycle


Can be recorded as an electrocardiogram (ECG or EKG)

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The control of heart rhythm


1 Pacemaker generates 2 Signals are delayed 3 Signals pass 4 Signals spread

wave of signals to contract.

at AV node.

to heart apex.

Throughout ventricles.

SA node (pacemaker)

AV node

Bundle branches Heart apex Purkinje fibers

ECG

Figure 42.8
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The pacemaker is influenced by


Nerves, hormones, body temperature, and exercise

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Concept 42.3: Physical principles govern blood circulation The same physical principles that govern the movement of water in plumbing systems
Also influence the functioning of animal circulatory systems

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Blood Vessel Structure and Function The infrastructure of the circulatory system
Is its network of blood vessels

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All blood vessels


Are built of similar tissues

Have three similar layers


Artery Vein Basement membrane Endothelium Endothelium Smooth muscle Connective tissue Artery

100 m
Valve Endothelium Smooth muscle Connective tissue Vein

Capillary

Venule

Figure 42.9

Arteriole

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Structural differences in arteries, veins, and capillaries


Correlate with their different functions

Arteries have thicker walls


To accommodate the high pressure of blood pumped from the heart

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In the thinner-walled veins


Blood flows back to the heart mainly as a result of muscle action
Direction of blood flow in vein (toward heart)
Valve (open)

Skeletal muscle

Valve (closed)

Figure 42.10
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Blood Flow Velocity Physical laws governing the movement of fluids through pipes
Influence blood flow and blood pressure

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Velocity (cm/sec) Pressure (mm Hg)

Area (cm2)

The velocity of blood flow varies in the circulatory system and is slowest in the capillary beds as a result of the high resistance and large total crosssectional area
5,000 4,000 3,000 2,000 1,000 0 50 40 30 20 10 0

120 100 80 60 40 20 0

Systolic pressure Diastolic pressure


Capillaries Venae cavae Arteries Venules Arterioles

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Veins

Aorta

Blood Pressure Blood pressure


Is the hydrostatic pressure that blood exerts against the wall of a vessel

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Systolic pressure
Is the pressure in the arteries during ventricular systole Is the highest pressure in the arteries

Diastolic pressure
Is the pressure in the arteries during diastole Is lower than systolic pressure

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Blood pressure
Can be easily measured in humans
1 A typical blood pressure reading for a 20-year-old is 120/70. The units for these numbers are mm of mercury (Hg); a blood pressure of 120 is a force that can support a column of mercury 120 mm high. 4 The cuff is loosened further until the blood flows freely through the artery and the sounds below the cuff disappear. The pressure at this point is the diastolic pressure remaining in the artery when the heart is relaxed. Blood pressure reading: 120/70 Pressure in cuff above 120 Rubber cuff inflated with air 120 Pressure in cuff below 120 120 70 Pressure in cuff below 70

Sounds audible in stethoscope

Sounds stop

Artery
Artery closed 2 A sphygmomanometer, an inflatable cuff attached to a pressure gauge, measures blood pressure in an artery. The cuff is wrapped around the upper arm and inflated until the pressure closes the artery, so that no blood flows past the cuff. When this occurs, the pressure exerted by the cuff exceeds the pressure in the artery. 3 A stethoscope is used to listen for sounds of blood flow below the cuff. If the artery is closed, there is no pulse below the cuff. The cuff is gradually deflated until blood begins to flow into the forearm, and sounds from blood pulsing into the artery below the cuff can be heard with the stethoscope. This occurs when the blood pressure is greater than the pressure exerted by the cuff. The pressure at this point is the systolic pressure.

Figure 42.12
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Blood pressure is determined partly by cardiac output


And partly by peripheral resistance due to variable constriction of the arterioles

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Capillary Function Capillaries in major organs are usually filled to capacity


But in many other sites, the blood supply varies

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Two mechanisms
Regulate the distribution of blood in capillary beds

In one mechanism
Contraction of the smooth muscle layer in the wall of an arteriole constricts the vessel

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In a second mechanism
Precapillary sphincters control the flow of blood between arterioles and venules
Precapillary sphincters Thoroughfare channel

(a) Sphincters relaxed Arteriole Venule Capillaries

(b) Sphincters contracted Arteriole Venule

(c) Capillaries and larger vessels (SEM)

Figure 42.13 ac
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20 m

The critical exchange of substances between the blood and interstitial fluid
Takes place across the thin endothelial walls of the capillaries

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The difference between blood pressure and osmotic pressure


Drives fluids out of capillaries at the arteriole end and into capillaries at the venule end
Tissue cell INTERSTITIAL FLUID Net fluid movement out Net fluid movement in

Capillary Capillary Red blood cell

15 m At the venule end of a capillary, blood pressure is less than osmotic pressure, and fluid flows from the interstitial fluid into the capillary. Blood pressure Osmotic pressure Inward flow

At the arterial end of a capillary, blood pressure is greater than osmotic pressure, and fluid flows out of the capillary into the interstitial fluid.

Direction of blood flow

Pressure

Outward flow

Figure 42.14

Arterial end of capillary

Venule end

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Fluid Return by the Lymphatic System The lymphatic system


Returns fluid to the body from the capillary beds Aids in body defense

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Fluid reenters the circulation


Directly at the venous end of the capillary bed and indirectly through the lymphatic system

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Concept 42.4: Blood is a connective tissue with cells suspended in plasma Blood in the circulatory systems of vertebrates
Is a specialized connective tissue

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Blood Composition and Function Blood consists of several kinds of cells


Suspended in a liquid matrix called plasma

The cellular elements


Occupy about 45% of the volume of blood

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Plasma Blood plasma is about 90% water Among its many solutes are
Inorganic salts in the form of dissolved ions, sometimes referred to as electrolytes

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The composition of mammalian plasma


Plasma 55%

Constituent
Water

Major functions Solvent for carrying other substances

Icons (blood electrolytes Sodium Potassium Calcium Magnesium Chloride Bicarbonate Plasma proteins Albumin Fibringen Immunoglobulins (antibodies)

Osmotic balance pH buffering, and regulation of membrane permeability

Separated blood elements

Osmotic balance, pH buffering Clotting Defense

Substances transported by blood Nutrients (such as glucose, fatty acids, vitamins) Waste products of metabolism Respiratory gases (O2 and CO2) Hormones
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Figure 42.15

Another important class of solutes is the plasma proteins


Which influence blood pH, osmotic pressure, and viscosity

Various types of plasma proteins


Function in lipid transport, immunity, and blood clotting

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Cellular Elements Suspended in blood plasma are two classes of cells


Red blood cells, which transport oxygen White blood cells, which function in defense

A third cellular element, platelets


Are fragments of cells that are involved in clotting

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The cellular elements of mammalian blood


Cellular elements 45%
Cell type Number per L (mm3) of blood 56 million Functions

Erythrocytes (red blood cells)

Transport oxygen and help transport carbon dioxide

Separated blood elements

Leukocytes (white blood cells)

5,00010,000

Defense and immunity

Basophil Eosinophil Neutrophil

Lymphocyte

Monocyte

Platelets

250,000 400,000

Blood clotting

Figure 42.15
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Erythrocytes Red blood cells, or erythrocytes


Are by far the most numerous blood cells

Transport oxygen throughout the body

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Leukocytes The blood contains five major types of white blood cells, or leukocytes
Monocytes, neutrophils, basophils, eosinophils, and lymphocytes, which function in defense by phagocytizing bacteria and debris or by producing antibodies

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Platelets Platelets function in blood clotting

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Stem Cells and the Replacement of Cellular Elements

The cellular elements of blood wear out


And are replaced constantly throughout a persons life

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Erythrocytes, leukocytes, and platelets all develop from a common source


A single population of cells called pluripotent stem cells in the red marrow of bones
Pluripotent stem cells (in bone marrow) Lymphoid stem cells Myeloid stem cells

Basophils B cells T cells

Lymphocytes

Eosinophils

Neutrophils Erythrocytes Platelets Monocytes

Figure 42.16
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Blood Clotting When the endothelium of a blood vessel is damaged


The clotting mechanism begins

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A cascade of complex reactions


Converts fibrinogen to fibrin, forming a clot
1 The clotting process begins when the endothelium of a vessel is damaged, exposing connective tissue in the vessel wall to blood. Platelets adhere to collagen fibers in the connective tissue and release a substance that makes nearby platelets sticky. 2 The platelets form a plug that provides emergency protection against blood loss. 3 This seal is reinforced by a clot of fibrin when vessel damage is severe. Fibrin is formed via a multistep process: Clotting factors released from the clumped platelets or damaged cells mix with clotting factors in the plasma, forming an activation cascade that converts a plasma protein called prothrombin to its active form, thrombin. Thrombin itself is an enzyme that catalyzes the final step of the clotting process, the conversion of fibrinogen to fibrin. The threads of fibrin become interwoven into a patch (see colorized SEM).

Collagen fibers Platelet releases chemicals that make nearby platelets sticky Platelet plug Fibrin clot Red blood cell

Clotting factors from: Platelets Damaged cells Plasma (factors include calcium, vitamin K)

Prothrombin

Thrombin Fibrinogen Fibrin


5 m

Figure 42.17

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Cardiovascular Disease Cardiovascular diseases


Are disorders of the heart and the blood vessels Account for more than half the deaths in the United States

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One type of cardiovascular disease, atherosclerosis


Is caused by the buildup of cholesterol within arteries
Connective tissue
Smooth muscle Endothelium Plaque

(a) Normal artery 50 m


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(b) Partly clogged artery

250 m

Figure 42.18a, b

Hypertension, or high blood pressure


Promotes atherosclerosis and increases the risk of heart attack and stroke

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 head
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Concept 42.5: Gas exchange occurs across specialized respiratory surfaces Gas exchange
Supplies oxygen for cellular respiration and disposes of carbon dioxide
Respiratory medium (air of water) Organismal level Circulatory system

O2

CO2

Respiratory surface

Cellular level Energy-rich molecules from food Cellular respiration ATP

Figure 42.19
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Animals require large, moist respiratory surfaces for the adequate diffusion of respiratory gases
Between their cells and the respiratory medium, either air or water

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Gills in Aquatic Animals Gills are outfoldings of the body surface


Specialized for gas exchange

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In some invertebrates
The gills have a simple shape and are distributed over much of the body
(a) Sea star. The gills of a sea star are simple tubular projections of the skin. The hollow core of each gill is an extension of the coelom (body cavity). Gas exchange occurs by diffusion across the gill surfaces, and fluid in the coelom circulates in and out of the gills, aiding gas transport. The surfaces of a sea stars tube feet also function in gas exchange.

Gills Coelom

Figure 42.20a
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Tube foot

Many segmented worms have flaplike gills


That extend from each segment of their body
(b) Marine worm. Many polychaetes (marine worms of the phylum Annelida) have a pair of flattened appendages called parapodia on each body segment. The parapodia serve as gills and also function in crawling and swimming.

Parapodia

Figure 42.20b
Gill

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The gills of clams, crayfish, and many other animals


Are restricted to a local body region
(c) Scallop. The gills of a scallop are long, flattened plates that project from the main body mass inside the hard shell. Cilia on the gills circulate water around the gill surfaces.

(d) Crayfish. Crayfish and other crustaceans have long, feathery gills covered by the exoskeleton. Specialized body appendages drive water over the gill surfaces.

Gills Gills

Figure 42.20c, d
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The effectiveness of gas exchange in some gills, including those of fishes


Is increased by ventilation and countercurrent flow of blood and water
Oxygen-poor blood Gill arch Blood vessel Oxygen-rich blood Lamella

Gill arch

Water flow

Operculum

O2 Water flow over lamellae showing % O2 Gill filaments

Figure 42.21

Blood flow through capillaries in lamellae showing % O2

Countercurrent exchange

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Tracheal Systems in Insects The tracheal system of insects


Consists of tiny branching tubes that penetrate the body
Air sacs Tracheae

Spiracle

(a) The respiratory system of an insect consists of branched internal tubes that deliver air directly to body cells. Rings of chitin reinforce the largest tubes, called tracheae, keeping them from collapsing. Enlarged portions of tracheae form air sacs near organs that require a large supply of oxygen. Air enters the tracheae through openings called spiracles on the insects body surface and passes into smaller tubes called tracheoles. The tracheoles are closed and contain fluid (blue-gray). When the animal is active and is using more O2, most of the fluid is withdrawn into the body. This increases the surface area of air in contact with cells. Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Figure 42.22a

The tracheal tubes


Supply O2 directly to body cells
Body cell Tracheole Air sac

Trachea Body wall Myofibrils

Air Tracheoles

Mitochondria

(b) This micrograph shows cross sections of tracheoles in a tiny piece of insect flight muscle (TEM). Each of the numerous mitochondria in the muscle cells lies within about 5 m of a tracheole.

Figure 42.22b
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2.5 m

Lungs Spiders, land snails, and most terrestrial vertebrates


Have internal lungs

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Mammalian Respiratory Systems: A Closer Look A system of branching ducts


Conveys air to the lungs
Branch from the pulmonary vein (oxygen-rich blood) Terminal bronchiole Nasal cavity Pharynx Larynx Esophagus Trachea Right lung Bronchus Bronchiole Diaphragm Heart SEM Left lung 50 m Branch from the pulmonary artery (oxygen-poor blood)

Alveoli

50 m

Colorized SEM

Figure 42.23
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In mammals, air inhaled through the nostrils


Passes through the pharynx into the trachea, bronchi, bronchioles, and dead-end alveoli, where gas exchange occurs

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Concept 42.6: Breathing ventilates the lungs The process that ventilates the lungs is breathing
The alternate inhalation and exhalation of air

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How an Amphibian Breathes An amphibian such as a frog


Ventilates its lungs by positive pressure breathing, which forces air down the trachea

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How a Mammal Breathes Mammals ventilate their lungs


By negative pressure breathing, which pulls air into the lungs
Rib cage expands as rib muscles contract Air inhaled Rib cage gets smaller as rib muscles relax Air exhaled

Lung Diaphragm

INHALATION Diaphragm contracts (moves down)

EXHALATION Diaphragm relaxes (moves up)

Figure 42.24
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Lung volume increases


As the rib muscles and diaphragm contract

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How a Bird Breathes Besides lungs, bird have eight or nine air sacs
That function as bellows that keep air flowing through the lungs
Air Anterior air sacs Trachea Posterior air sacs Lungs Lungs Air

Air tubes (parabronchi) in lung

1 mm

INHALATION Air sacs fill

EXHALATION Air sacs empty; lungs fill

Figure 42.25
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Air passes through the lungs


In one direction only

Every exhalation
Completely renews the air in the lungs

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Control of Breathing in Humans The main breathing control centers


Are located in two regions of the brain, the medulla oblongata and the pons
1 The control center in the medulla sets the basic rhythm, and a control center in the pons moderates it, smoothing out the transitions between inhalations and exhalations. Cerebrospinal fluid 4 The medullas control center also helps regulate blood CO2 level. Sensors in the medulla detect changes in the pH (reflecting CO2 concentration) of the blood and cerebrospinal fluid bathing the surface of the brain. 5 Nerve impulses relay changes in CO2 and O2 concentrations. Other sensors in the walls of the aorta and carotid arteries in the neck detect changes in blood pH and send nerve impulses to the medulla. In response, the medullas breathing control center alters the rate and depth of breathing, increasing both to dispose of excess CO2 or decreasing both if CO2 levels are depressed.

Pons 2 Nerve impulses trigger muscle contraction. Nerves from a breathing control center in the medulla oblongata of the brain send impulses to the diaphragm and rib muscles, stimulating them to contract and causing inhalation. Breathing control centers

Medulla oblongata

Carotid arteries Aorta 3 In a person at rest, these nerve impulses result in about 10 to 14 inhalations per minute. Between inhalations, the muscles relax and the person exhales.

Figure 42.26

Diaphragm Rib muscles

6 The sensors in the aorta and carotid arteries also detect changes in O2 levels in the blood and signal the medulla to increase the breathing rate when levels become very low.

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The centers in the medulla


Regulate the rate and depth of breathing in response to pH changes in the cerebrospinal fluid

The medulla adjusts breathing rate and depth


To match metabolic demands

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Sensors in the aorta and carotid arteries


Monitor O2 and CO2 concentrations in the blood Exert secondary control over breathing

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Concept 42.7: Respiratory pigments bind and transport gases The metabolic demands of many organisms
Require that the blood transport large quantities of O2 and CO2

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The Role of Partial Pressure Gradients Gases diffuse down pressure gradients
In the lungs and other organs

Diffusion of a gas
Depends on differences in a quantity called partial pressure

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A gas always diffuses from a region of higher partial pressure


To a region of lower partial pressure

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In the lungs and in the tissues


O2 and CO2 diffuse from where their partial pressures are higher to where they are lower

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Inhaled air

Exhaled air

160 0.2 O2 CO2

Alveolar spaces
104 40

120 27

O2 CO2 O2 CO2 CO2 O2

Alveolar epithelial cells


Blood entering alveolar capillaries
40 45

1
O2

Blood leaving alveolar capillaries


104 40

O2 CO2

Alveolar capillaries of lung

O2 CO2

Pulmonary arteries

Pulmonary veins

Systemic veins
CO2

Systemic arteries
Heart Tissue capillaries
O2

Blood leaving tissue capillaries


40 45

Blood entering tissue capillaries


100 40

CO2

O2

O2 CO2

O2 CO2

Tissue cells

Figure 42.27
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<40 >45

O2 CO2

Respiratory Pigments Respiratory pigments


Are proteins that transport oxygen

Greatly increase the amount of oxygen that blood can carry

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Oxygen Transport The respiratory pigment of almost all vertebrates


Is the protein hemoglobin, contained in the erythrocytes

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Like all respiratory pigments


Hemoglobin must reversibly bind O2, loading O2 in the lungs and unloading it in other parts of the body
Heme group
Iron atom

O2 loaded in lungs O2 unloaded In tissues

O2

O2

Figure 42.28

Polypeptide chain

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Loading and unloading of O2


Depend on cooperation between the subunits of the hemoglobin molecule

The binding of O2 to one subunit induces the other subunits to bind O2 with more affinity

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Cooperative O2 binding and release


Is evident in the dissociation curve for hemoglobin

A drop in pH
Lowers the affinity of hemoglobin for O2

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O2 saturation of hemoglobin (%)

(a) PO2 and Hemoglobin Dissociation at 37C and pH 7.4

100 80 60 40 20 0 0

O2 unloaded from hemoglobin during normal metabolism O2 reserve that can be unloaded from hemoglobin to tissues with high metabolism

20

40

60

80 100
Lungs

Tissues during Tissues at rest exercise PO2 (mm Hg) (b) pH and Hemoglobin Dissociation

O2 saturation of hemoglobin (%)

100 80 60 40 20 0 0 20 40
pH 7.2 pH 7.4
Bohr shift: Additional O2 released from hemoglobin at lower pH (higher CO2 concentration)

Figure 42.29a, b
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60

80 100

PO2 (mm Hg)

Carbon Dioxide Transport Hemoglobin also helps transport CO2


And assists in buffering

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Carbon from respiring cells


Diffuses into the blood plasma and then into erythrocytes and is ultimately released in the lungs

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Carbon dioxide produced by body tissues diffuses into the interstitial fluid and the plasma. Over 90% of the CO2 diffuses into red blood cells, leaving only 7% in the plasma as dissolved CO2. Some CO2 is picked up and transported by hemoglobin.

Tissue cell CO2 produced Interstitial CO 2 fluid 1 Blood plasma CO 2 within capillary 2 CO2 H2O 4 3

CO2 transport from tissues

Most of the HCO3 diffuse into the plasma where it is carried in the bloodstream to the lungs. In the HCO3 diffuse from the plasma red blood cells, combining with H+ released from hemoglobin and forming H2CO3. Carbonic acid is converted back into CO2 and water. CO2 formed from H2CO3 is unloaded from hemoglobin and diffuses into the interstitial fluid. diffuses into the alveolar space, from which it is expelled during exhalation. The reduction of CO2 concentration in the plasma drives the breakdown of H2CO3 Into CO2 and water in the red blood cells (see step 9), a reversal of the reaction that occurs in the tissues (see step 4).

Capillary wall

Red Hemoglobin H2CO3 blood Carbonic acid Hb picks up cell CO2 and H+ 5 + H+ Bicarbonate
HCO3

However, most CO2 reacts with water in red blood cells, forming carbonic acid (H2CO3), a reaction catalyzed by carbonic anhydrase contained. Within red blood cells. Carbonic acid dissociates into a biocarbonate ion (HCO3) and a hydrogen ion (H+).

HCO3

To lungs

10
HCO3
CO2 transport to lungs

8
HCO3 + H+

11 CO2
Hemoglobin releases CO2 and H+

H2CO3

Hb

9 H2O CO2

Hemoglobin binds most of the H+ from H2CO3 preventing the H+ from acidifying the blood and thus preventing the Bohr shift.

CO2 CO2 10 CO2 11 Alveolar space in lung

Figure 42.30

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Elite Animal Athletes Migratory and diving mammals


Have evolutionary adaptations that allow them to perform extraordinary feats

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The Ultimate Endurance Runner The extreme O2 consumption of the antelopelike pronghorn
Underlies its ability to run at high speed over long distances

Figure 42.31
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Diving Mammals Deep-diving air breathers


Stockpile O2 and deplete it slowly

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