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BIOLOGY

Animal Transport
Systems

彭志維老師
北醫生物學工程學系
Overview:Trading Places
• Every organism must exchange materials with its
environment
• Exchanges ultimately occur at the cellular level by crossing
the plasma membrane
• 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
• Internal transport and gas exchange are functionally related in most
animals
• Gills (鰓) are an example of a specialized exchange system in animals
• O2 diffuses from the water into blood vessels
• CO2 diffuses from blood into the water 2
Section I: Circulatory systems link
exchange surfaces with cells
throughout the body
• In small and/or thin animals, cells can exchange materials
directly by diffusion (擴散) with the surrounding medium
• Diffusion time is proportional to the square of the distance
• Diffusion is only efficient over small distances
• In most animals, cells exchange materials with the
environment via a fluid-filled circulatory system
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Gastrovascular Cavities
• Some animals lack a circulatory system
• Some cnidarians (刺胞動物), such as jellies, have
elaborate gastrovascular cavities (消化循環腔)
• A gastrovascular cavity functions in both digestion and
distribution of substances throughout the body
• The body wall that encloses the gastro-
vascular cavity is only two cells thick
• Flatworms have a gastrovascular
cavity and a large surface area to
volume ratio
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Evolutionary Variation in
Circulatory Systems
• A circulatory system minimizes the diffusion distance in
animals with many cell layers.
• A circulatory system has
• A circulatory fluid (循環液體)
• A set of interconnecting vessels
• A muscular pump (唧筒), the heart
• The system connects the fluid that surrounds cells with the organs
that exchange gases, absorb nutrients, and dispose of wastes.
• Circulatory systems can be open or closed, and vary in the number of
circuits in the body.
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Open and Closed
Circulatory Systems
• In insects, other arthropods(節肢動物), and most
molluscs ( 軟 體 動 物 ), 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 called
hemolymph (血淋巴).
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• In a closed circulatory system, blood is confined to
vessels and distinct from the interstitial fluid
• Closed systems are more efficient at transporting
circulatory fluids to tissues and cells
• Annelids (環節動物), cephalopods (頭足動物), and
vertebrates have closed circulatory systems
Organization of Vertebrate Circulatory Systems
• Humans and other vertebrates have a closed circulatory
system called the cardiovascular system (心血管系統)
• The three main types of blood vessels are arteries, veins, and
capillaries
• Blood flow is one way in these vessels
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• Arteries (動脈) branch into arterioles (小動脈) and
carry blood away from heart to capillaries (微血管).
• Networks of capillaries called capillary beds (微血
管網), are the sites of chemical exchange
between the blood and interstitial fluid
• Venules (小靜脈) converge into veins (靜脈), and return blood
from capillaries to the heart
• Arteries and veins are distinguished by the direction of blood
flow, not by O2 content
• Vertebrate hearts contain two or more chambers
• Blood enters through an atrium (心房) and is pumped out
through a ventricle (心室). 8
Single Circulation (單循環)
• Bony fishes(硬骨魚), rays (魟), and sharks have
single circulation with a two-chambered heart
• In single circulation, blood leaving
the heart passes through two
capillary beds before returning

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Double Circulation (雙循環)
• Amphibian (兩棲類), reptiles (爬蟲類), and mammals
have double circulation
• Oxygen-poor and oxygen-rich blood are pumped
separately from the right and left sides of the heart
• In reptiles and mammals, oxygen-poor blood flows through the
pulmonary circuit (肺循環) to pick up oxygen through the lungs
• In amphibians, oxygen-poor blood flows through a
pulmocutaneous circuit (肺皮循環) to pick up oxygen through
the lungs and skin
• Oxygen-rich blood delivers oxygen through the systemic circuit
(體循環)
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Evolutionary Variation in
Double Circulatory
• Hearts vary in different vertebrate groups
• Double circulation maintains
higher blood pressure in
organs than does single
circulation
• Amphibians and many reptiles
may pass long periods without
gas exchange or relying on
gas exchange from another
tissue, usually the skin. 11
肺皮循環
Amphibians
• Frogs and other amphibians have a
three-chambered heart:two atria and
one ventricle
• The ventricle pumps blood into a
forked artery that splits the ventricle’s
output into the pulmocutaneous circuit
and the systemic circuit
體循環 • When underwater, blood flow to the
lungs is nearly shut off 12
Reptiles (Except Birds)

(肺循環)
• Turtles, snakes, and lizards (蜥蜴) have
a three-chambered heart:two atria
and one ventricle
• In alligators(鱷) septum divides the
ventricle
• Reptiles have double circulation, with
a pulmonary circuit (lungs) and a
(體循環)
systemic circuit
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Mammals and Birds
• Have four-chambered heart with
two atria and two ventricles
• Left side of heart pumps and
receives only oxygen-rich blood,
while the right side receives and
pumps only oxygen-poor blood
• Mammals and birds are
endotherms(內溫動物)
and require more O2 than
ectotherms(外溫動物)
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Section (II):
Mammalian Circulation
• The mammalian cardiovascular system meets the body’s
continuous demand for O2
• Blood begins its flow with the right ventricle pumping blood to lungs
• In the lungs, the blood loads O2 and unloads CO2
• Oxygen-rich blood from lungs enters the heart at the left atrium and is
pumped through the aorta (主動脈) to body tissues by the left ventricle
• The aorta provides blood to heart through coronary arteries (冠狀動脈)
• Blood returns to the heart through the superior vena cava上腔大靜脈
(blood from head, neck, and forelimbs) and inferior vena cava下腔大靜
脈(blood from trunk and hind limbs)
• The superior vena cava and inferior vena cava flow into the right atrium
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循環起始點

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The Mammalian Heart:
A Closer Look
• A closer look at the mammalian heart provides a
better understanding of double circulation

半月辦

房室辦
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• The heart rate, also called the pulse, is the
number of beats per minute
• The stroke volume (心搏量) is the amount of blood
pumped in a single contraction
• The cardiac output (心輸出量) 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 (房室辦) separate each
atrium and ventricle
• The semilunar valves (半月辦) control blood flow to the
aorta and the pulmonary artery
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心房收縮期/心室舒張期
• The heart contracts and
relaxes in a rhythmic
cycle called cardiac
心房/心室舒張期
cycle(心週期)
• The contraction, or pumping,
phase is called systole(心縮期)
• The relaxation, or filling, phase is
called diastole(心舒期)
• 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
心室收縮期/心房舒張期
murmur(心雜音) 19
Maintaining the Heart’s
Rhythmic Beat
• Some cardiac muscle cells are self-excitable, meaning they
contract without any signal from the nervous system
• The sinoatrial (SA) node (竇房結), or pacemaker (節律點), sets the rate
and timing at which cardiac muscle cells contract
• Impulses from the SA node travel to the atrioventricular (AV) node
(房室結)
• At the AV node, the impulses are delayed and then travel to the
Purkinje fibers (浦金氏纖維) that make the ventricles contract
• Impulses that travel during the cardiac cycle can be recorded as an
electrocardiogram (ECG or EKG) (心電圖)
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• The pacemaker is regulated by two portions of the
nervous system: sympathetic (交感) and
parasympathetic (副交感) divisions
• The sympathetic division speeds up the pacemaker
• The parasympathetic division slows down the pacemaker
• The pacemaker is also regulated by hormones and temperature

竇房節 特化
房室結 浦金氏纖維
心肌纖維
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Section (III): Blood Vessel Structure and
Function
• A vessel’s cavity is called central lumen (中央內腔).
• The epithelial layer that lines blood vessels is
called the endothelium(內皮)
• The endothelium is smooth and minimizes resistance
• Capillaries have thin walls, the endothelium plus its basal
lamina (基底膜), 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 22
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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 (總橫截面積)
• Blood flow in capillaries is necessarily
slow for exchange of materials
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Blood Pressure
• Blood flows from areas of higher pressure to areas of
lower pressure
• Blood pressure is the 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(收縮壓) is the pressure in the arteries during
ventricular systole; it is the highest pressure in the arteries
• Diastolic pressure (舒張壓) is the pressure in the arteries during
diastole; it is lower than systolic pressure
• A pulse(脈搏) is the rhythmic bulging of artery walls with each
heartbeat 25
Blood pressure reading: 120/70

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Regulation of Blood Pressure
• Blood pressure is determined by cardiac output
(心輸出量) and peripheral resistance (周邊阻力)
due to constriction of arterioles
• Vasoconstriction (血管 收縮) is the contraction of smooth
muscle in arteriole walls; it increases blood pressure
• Vasodilation (血管舒張) 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
• Nitric oxide (一氧化氮) is a major inducer of vasodilation
• The peptide endothelin(內皮素) is an important inducer of
vasoconstriction 27
Blood Pressure and Gravity
• 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 and 70 mm Hg at diastole
• Fainting (昏暈) is caused by inadequate blood flow to the head
• Animals with longer necks require a higher
systolicpressure 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 (瓣膜) in veins prevent
backflow of blood 28
Capillary Function
• Blood flows through only 510% of the body’s
capillaries at a time.
• Blood supply varies in many other sites.
• Capillaries in major organs are usually filled to capacity
• 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 (微血管前括約肌) control flow of
blood between arterioles and venules
• Blood flow is regulated by nerve impulses, hormones, and other
chemicals
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• The exchange of substances between the blood
and interstitial fluid takes place across the thin
endothelial walls of capillaries
• The difference between blood pressure (血壓)
and osmotic pressure (滲透壓) drives fluids out of
capillaries at the arteriole end and into
capillaries at the venule end
• Most blood proteins and all
blood cells are too large to
pass through the endothelium
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Fluid Return by the Lymphatic
System
• The lymphatic system (淋巴系統) returns fluid that
leaks out from the capillary beds
• Fluid, called lymph(淋巴) , reenters the circulation directly
at the venous end of the capillary bed and indirectly
through the lymphatic system
• The lymphatic system drains into veins in the neck
• Valves in lymph vessels prevent the backflow of fluid
• Lymph nodes (淋巴結) are organs that filter lymph and
play an important role in the body’s defense
• Edema (水腫) is swelling caused by disruptions in the flow
of lymph
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Section (IV):
Blood Composition and Function
• Blood consists of several kinds of cells suspended in a
liquid matrix called plasma (血漿) (55%); The cellular elements
occupy about 45% of the volume of blood
• Blood plasma is about 90% water
• Among its solutes are inorganic salts in the form of dissolved
ions, sometimes called electrolytes(電解質), and the plasma
proteins ( 血 漿 蛋 白 ), which influence blood pH, osmotic
pressure, and viscosity
• Various plasma proteins function in lipid transport, immunity,
and blood clotting 34
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Cellular Elements
• Suspended in blood plasma are two types of cells
 Red blood cells(紅血球;erythrocytes) transport oxygen O2
 White blood cells (白血球;leukocytes) function in defense
• Platelets(血小板) , a third cellular element, are fragments of
cells that are involved in clotting

Erythrocytes
• Red blood cells are by far the most numerous blood cells
• They contain hemoglobin(血紅素), the iron-containing protein that
transports O2
• Each molecule of hemoglobin binds up to four molecules of O2
• In mammal, mature erythrocytes lack nuclei and mitochondria
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• Sickle-cell disease(鎌刀型貧血症) is caused by abnormal
hemoglobin proteins that form aggregates
• The aggregates can deform an erythrocyte into a sickle
shape
• Sickled cells can rupture, or block blood vessels
Leukocytes
• There are five major types of white blood cells: monocytes (單
核球),neutrophils(嗜中性球), basophils(嗜鹼性球),
eosinophils(嗜酸性球), and lymphocytes(淋巴球)
• They function in defense by phagocytizing bacteria and debris
or by producing antibodies
• They are found both in and outside of the circulatory system
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Platelets (血小板)
• Platelets are fragments of cells and function in
blood clotting
• Coagulation is the formation of a solid clot from liquid
blood
• A cascade of complex reactions converts inactive
fibrinogen (血纖蛋白原) to fibrin (血纖蛋白), forming a clot
• A blood clot formed within a blood vessel is called a
thrombus and can block blood flow
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Stem Cells and the Replacement
of Cellular Elements
• The cellular elements of blood wear out
and are being replaced constantly
• Erythrocytes, leukocytes, and platelets 骨髓
all develop from a common source of
stem cells (幹細胞) in the red marrow of
淋巴前驅細胞 骨髓前驅細胞
bones, especially ribs, vertebrae,
sternum, and pelvis
• The hormone erythropoietin (紅血球生成
素;EPO) stimulates erythrocyte
production when O2 delivery is low 41
Cardiovascular Disease
(心血管疾病)
• Cardiovascular diseases are disorders of the heart and
the blood vessels
• Cardiovascular diseases account for more than half the deaths
in the United States
• Cholesterol (膽固醇), a steroid, helps maintain membrane
fluidity
• Low-density lipoprotein (LDL; 低密度脂蛋白) delivers
cholesterol to cells for membrane production
• High-density lipoprotein (HDL; 高密度脂蛋白) scavenges
cholesterol for return to the liver
• Risk for heart disease increases with a high LDL to HDL ratio
• Inflammation is also a factor in cardiovascular disease
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Atherosclerosis, Heart Attacks,
and Stroke
• One type of
cardiovascular disease,
atherosclerosis(粥狀動脈
硬化), is caused by the
buildup of plaque
deposits within arteries

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44
• A heart attack, or myocardial infarction (心肌梗塞),
is the death of cardiac muscle tissue resulting
from blockage of one or more coronary arteries
• Coronary arteries(冠狀動脈)supply oxygen-rich blood to the
heart muscle
• A stroke (中風) is the death of nervous tissue in the brain,
usually resulting from rupture or blockage of arteries in the
head
• Angina pectoris(心絞痛) is caused by partial blockage of
the coronary arteries and results in chest pains
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Risk Factors and Treatment of
Cardiovascular Disease
• A high LDL to HDL ratio increases the risk of cardiovascular
disease
• The proportion of LDL relative to HDL can be decreased by exercise,
not smoking, and avoiding foods with trans fats (反式脂肪)
• Drugs called statins (降膽固醇類藥物) reduce LDL levels and risk of
heart attacks
• Inflammation plays a role in atherosclerosis and thrombus
formation. Aspirin inhibits inflammation and reduces the risk of
heart attacks and stroke
• Hypertension ( 高 血 壓 ), or 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 46
Section (V): Gas exchange occurs across
specialized respiratory surfaces
• Gas exchange(氣體交換) supplies O2 for cellular
respiration and disposes of CO2
• Partial pressure (分壓)is the pressure exerted by a particular gas
in a mixture of gases. A gas diffuses from a region of higher
partial pressure to a region of lower partial pressure
• Gases diffuse down pressure gradients in the lungs and other
organs as a result of differences in partial pressure
• Animals can use air or water as a source of O2, or respiratory
medium (呼吸介質)
• In a given volume, there is less O2 available in water than in air
Obtaining O2 from water requires greater efficiency than air
breathing 47
Respiratory Surfaces
• Animals require large, moist respiratory surfaces
(呼吸表面) for exchange of gases between their cells and
the respiratory medium, either air or water
• Gas exchange across respiratory surfaces takes place by
diffusion
• Respiratory surfaces vary by animal and can include the
outer surface, skin (皮膚), gills (鰓), tracheae (氣管), and
lungs (肺) 48
Gills (鰓) in Aquatic Animals
• Gills are outfoldings of the body that create a
large surface area for gas exchange
• Ventilation (換氣) moves the respiratory medium over the
respiratory surface
• Aquatic animals move through water or move water over
their gills for ventilation
• Fish gills use a countercurrent exchange (逆流交換),
system, where blood flows in the opposite direction to
water passing over the gills; blood is always less saturated
with O2 than the water it meets 49
體腔

疣足
管足
海蟲 螯蝦 海星 50
鰓弧

鰓板
鰓蓋

逆流交換

鰓絲

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Tracheal Systems in
Insects
• The tracheal system (氣管系統) of insects consists of tiny
branching tubes that penetrate the body
• The tracheal tubes supply O2 directly to body cells
• The respiratory and circulatory systems are separate
• Larger insects must ventilate their tracheal system to meet
O2 demands
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氣管
氣囊

氣門
53
Lungs
• Lungs are an infolding of the body surface
• The circulatory system (open or closed) transports
gases between the lungs and the rest of the body
• The size and complexity of lungs correlate with an animal’s
metabolic rate
• A system of branching ducts conveys air to the lungs
• Air inhaled through the nostrils is warmed, humidified, and
sampled for odors
• The pharynx (咽)directs air to the lungs and food to the
stomach
• Swallowing tips the epiglottis (會厭) over the glottis(聲門) in
the pharynx to prevent food from entering the trachea 54
• Air passes through the pharynx (咽), larynx (喉),
trachea (氣管), bronchi (支氣管), and bronchioles
(小支氣管) to the alveoli (肺泡), where gas exchange
occurs
• Exhaled air passes over the vocal cords (聲帶) in the
larynx to create sounds
• Cilia (纖毛)and mucus(黏液) line the epithelium of the air ducts
and move particles up to the pharynx
• This “mucus escalator” cleans the respiratory system and
allows particles to be swallowed into the esophagus(食道)
• Gas exchange takes place in alveoli, air sacs at the tips of
bronchioles
• Oxygen diffuses through the moist film of the epithelium and
into capillaries
• Carbon dioxide diffuses from the capillaries across the
epithelium and into the air space 55
終端
小支氣管

鼻腔

肺泡
氣管

支氣管
小支氣管
橫膈
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• Alveoli lack cilia and are susceptible to contamination
• Secretions called surfactants(介面活性劑) coat the
surface of the alveoli
• Preterm babies(早產兒) lack surfactant and are
vulnerable to respiratory distress syndrome (呼吸窘迫
症); treatment is provided by artificial surfactants
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Section (VI): Breathing ventilates
the lungs
• The process that ventilates the lungs is breathing, the
alternate inhalation and exhalation of air
How an Amphibian Breathes
• An amphibian such as a frog ventilates its lungs by positive
pressure breathing(正壓呼吸), which forces air down the trachea
How a Bird Breathes
• Birds have eight or nine air sacs (氣囊) that function as bellows
that keep air flowing through the lungs
• Air passes through the lungs in one direction only
• Every exhalation completely renews the air in the lungs 58
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How a Mammal Breathes
• Mammals ventilate their lungs by negative
pressure breathing, which pulls air into the lungs
• Lung volume increases as the rib muscles (肋間肌) and
diaphragm (橫膈) contract
• The tidal volume(潮氣容積) is the volume of air inhaled
with each breath
• The maximum tidal volume is the vital capacity (肺活量)
• After exhalation, a residual volume (肺餘容積) of air
remains in the lungs
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Control of Breathing in
Humans
• In humans, the main breathing control centers
(呼吸控制中心) are in two regions of the brain, the
medulla oblongata (延腦) and the pons (橋腦)
• The medulla regulates the rate and depth of breathing in
response to pH changes in the cerebrospinal fluid (腦脊髓液)
• The medulla adjusts breathing rate and depth to match
metabolic demands
• The pons regulates the tempo
• Sensors in the aorta (大動脈) and carotid arteries (頸動脈)
monitor O2 and CO2 concentrations in the blood
• These sensors exert secondary control over breathing 62
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Section (VII): 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
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Respiratory Pigments
• Respiratory pigments(呼吸色素), proteins that
transport oxygen, greatly increase the amount of
oxygen that blood can carry
• Arthropods (節肢動物) and many molluscs (軟體動物) have
hemocyanin (血青素) with copper as the oxygen-binding
component
• Most vertebrates and some invertebrates use hemoglobin (血紅
素)
• In vertebrates, hemoglobin is contained within erythrocytes
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Hemoglobin(血紅素)
• A single hemoglobin molecule can
carry four molecules of O2, one
molecule for each iron containing
heme(血基質) group
• The hemoglobin dissociation curve (解
離曲線) 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(波爾
移轉效應) 67
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Carbon Dioxide Transport
CO2 from respiring cells diffuses into the blood and
is transported either in blood plasma, bound to hemoglobin,
or as bicarbonate ions (HCO3–,碳酸根)
Only about 7% of CO2 from respiring cells diffuses into the
blood and is transported in blood plasma, bound to
hemoglobin
The remainder reacts with water to form H2CO3, which
dissociates into H+ and bicarbonate ions (HCO3–)
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Respiratory Adaptations
of Diving Mammals
• Diving mammals (潛泳動物) have evolutionary adaptations
that allow them to perform extraordinary feats
• For example, Weddell seals in Antarctica (南極洲海豹) can
remain underwater for 20 minutes to an hour
• For example, elephant seals (象鼻海豹) can dive to 1,500
m and remain underwater for 2 hours
• These animals have a high blood to body volume ratio
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• Deep-diving air breathers stockpile O2 and deplete it slowly
• Diving mammals can store oxygen in their muscles in
myoglobin (肌紅素) proteins
• Diving mammals also conserve oxygen by
• Changing their buoyancy (浮力)to glide passively
• Decreasing blood supply to muscles
• Deriving ATP in muscles from fermentation once
oxygen is depleted
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