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Apago PDF Enhancer I Chapter 50 The Circulatory System Chapter Outline 50.1 The Components of Blood 50.2 Invertebrate Circulatory Systems 50.3 Vertebrate Circulatory Systems 50.4 The Four-Chambered Heart and the Blood Vessels 50.5 Characteristics of Blood Vessels 50.6 Regulation of Blood Flow and Blood Pressure Introduction In multicellular organisms, oxygen obtained by the respiratory system and nutrients processed by the digestive system must be transported to cells throughout the body. Conversely, carbon dioxide and other waste products produced within the cells must be returned to the respiratory, digestive, and urinary systems for elimination from the body. These tasks are the responsibility of the circulatory system. All multicellular organisms have a heart that pumps fluids through the body. Many invertebrates have an open system in which fluids move through the body cavity. Vertebrates also have a system like this that moves lymph through the body; however, the primary circulatory fluid is blood, which means through a closed system of blood vessels. platelets, although included in figure 50.1, are not complete cells; rather, they are fragments of cells that are produced in the bone marrow. (We describe the action of platelets in blood clotting later in this section.) Circulating blood has many functions: Transportation. 1. All of the substances essential for cellular metabolism are transported by blood. Red blood cells transport oxygen attached to hemoglobin; nutrient molecules are carried in the plasma, sometimes bound to carriers; and metabolic wastes are eliminated as blood passes through the liver and kidneys. Regulation. 2. The cardiovascular system transports regulatory hormones from the endocrine glands and also 50.1 The Components of Blood Learning Outcomes Describe the functions of circulating blood. 1. Distinguish between the types of formed elements. 2. Delineate the process of blood clotting. 3. Blood is a connective tissue composed of a fluid matrix, called plasma, and several different kinds of cells and other formed elements that circulate within that fluid (figure 50.1). Blood CHAPTER Apago PDF Enhancer Figure 50.1 Composition of blood. participates in temperature regulation. Contraction and dilation of blood vessels near the surface of the body, beneath the epidermis, helps to conserve or to dissipate heat as needed. Protection. 3. The circulatory system protects against injury and foreign microbes or toxins introduced into the body. Blood clotting helps to prevent blood loss when vessels are damaged. White blood cells, or leukocytes, help to disarm or disable invaders such as viruses and bacteria (see chapter 52). Blood plasma is a fluid matrix Blood plasma is the matrix in which blood cells and platelets are suspended. Interstitial (extracellular) fluids originate from the fluid present in plasma. Although plasma is 92% water, it also contains the follow- ing solutes: Nutrients, wastes, and hormones. 1. Dissolved within the plasma are all of the nutrients resulting from digestive breakdown that can be used by cells, including glucose, amino acids, and vitamins. Also dissolved in the plasma are wastes such as nitrogen compounds and CO 2 produced by metabolizing cells. Endocrine hormones released from glands are also carried through the blood to their target cells. Ions. 2. Blood plasma is a dilute salt solution. The predominant plasma ions are Na + , Cl , and bicarbonate ions (HCO 3 ). In addition, plasma contains trace amounts of other ions such as Ca 2+ , Mg 2+ , Cu 2+ , K + , and Zn 2+ . Proteins. 3. As mentioned earlier, the liver produces most of the plasma proteins, including albumin, which constitutes most of the plasma protein; the alpha (α) and beta (β) globulins, which serve as carriers of lipids and steroid hormones; and fibrinogen, which is required for blood clotting. Blood plasma with the fibrinogen removed is called serum. Formed elements include circulating cells and platelets The formed elements of blood cells and cell fragments include red blood cells, white blood cells, and platelets. Each element has a specific function in maintaining the body’s health and homeostasis. Erythrocytes Each microliter of blood contains about 5 million red blood cells, or erythrocytes. The fraction of the total blood volume that is occupied by erythrocytes is called the blood’s hematocrit; in humans, the hematocrit is typically around 45%. Each erythrocyte resembles a doughnut-shaped disk with a central depression that does not go all the way through. Ma- ture mammalian erythrocytes lack nuclei. The erythrocytes of vertebrates contain hemoglobin, a pigment that binds and transports oxygen. (Hemoglobin was described more fully in the previous chapter when we discussed respiration.) In verte- brates, hemoglobin is found only in erythrocytes. In inverte- brates, the oxygen-binding pigment (not always hemoglobin) is also present in plasma. Leukocytes Less than 1% of the cells in human blood are white blood cells, or leukocytes; there are only 1 or 2 leukocytes for every 1000 erythrocytes. Leukocytes are larger than erythrocytes and have nuclei. Furthermore, leukocytes are not confined to the blood as erythrocytes are, but can migrate out of capillaries through the intercellular spaces into the surrounding intersti- tial (tissue) fluid. Leukocytes come in several varieties, each of which plays a specific role in defending against invading microorganisms and other foreign substances, as described in chapter 52. Granular leukocytes include neutrophils, eosinophils, and ba- sophils, which are named according to the staining properties of granules in their cytoplasm. Nongranular leukocytes in- clude monocytes and lymphocytes. In humans, neutrophils are the most numerous of the leukocytes, followed in order by lym- phocytes, monocytes, eosinophils, and basophils. chapter 50 The Circulatory System 1019 www.ravenbiology.com

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    I

    Ch

    ap

    ter 50

    The

    Cir

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    cie

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    ula

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    lop

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    rre

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    y w

    ith

    gil

    ls

    Ch

    ord

    ates

    an

    cest

    ral

    to t

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    ebra

    tes

    are

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    to h

    ave

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    si

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    eart

    s, s

    imil

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    o t

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    ow

    see

    n in

    lan

    cele

    ts (

    see

    chap

    ter

    35

    ). T

    he

    hea

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    ore

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    an a

    sp

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    f th

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    teri

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    on

    trac

    ted

    in

    sim

    ple

    per

    ista

    ltic

    wav

    es.

    T

    he

    dev

    elo

    pm

    ent

    of

    gil

    ls b

    y fi

    shes

    req

    uir

    ed a

    mo

    re e

    ffi-

    cien

    t p

    um

    p,

    and

    in

    fi

    shes

    w

    e se

    e th

    e ev

    olu

    tio

    n

    of

    a tr

    ue

    cham

    ber

    -pu

    mp

    hea

    rt. T

    he

    fish

    hea

    rt i

    s, i

    n e

    ssen

    ce, a

    tu

    be

    wit

    h

    fou

    r st

    ruct

    ure

    s ar

    raye

    d o

    ne

    afte

    r th

    e o

    ther

    to

    fo

    rm t

    wo

    pu

    mp

    -in

    g c

    ham

    ber

    s (f

    igu

    re 5

    0.5

    ) . T

    he

    firs

    t tw

    o s

    tru

    ctu

    res

    the

    sin

    us

    ven

    osu

    s an

    d a

    triu

    m

    form

    th

    e fi

    rst

    cham

    ber

    ; th

    e se

    con

    d t

    wo

    , th

    e ve

    ntr

    icle

    an

    d c

    on

    us

    arte

    rio

    sus,

    fo

    rm t

    he

    seco

    nd

    ch

    amb

    er.

    Th

    e si

    nu

    s ve

    no

    sus

    is t

    he

    firs

    t to

    co

    ntr

    act,

    foll

    ow

    ed b

    y th

    e at

    riu

    m,

    the

    ven

    tric

    le, a

    nd

    fin

    ally

    th

    e co

    nu

    s ar

    teri

    osu

    s.

    Des

    pit

    e sh

    ifts

    in

    th

    e re

    lati

    ve p

    osi

    tio

    ns

    of

    thes

    e st

    ruct

    ure

    s,

    this

    hea

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    eat

    seq

    uen

    ce i

    s m

    ain

    tain

    ed i

    n a

    ll v

    erte

    bra

    tes.

    In

    fis

    h,

    the

    elec

    tric

    al i

    mp

    uls

    e th

    at p

    rod

    uce

    s th

    e co

    ntr

    acti

    on

    is

    init

    iate

    d

    in t

    he

    sin

    us

    ven

    osu

    s; in

    oth

    er v

    erte

    bra

    tes,

    th

    e el

    ectr

    ical

    imp

    uls

    e is

    in

    itia

    ted

    by

    a st

    ruct

    ure

    ho

    mo

    log

    ou

    s to

    th

    e si

    nu

    s ve

    no

    sus

    the

    sin

    oat

    rial

    (S

    A)

    no

    de.

    A

    fter

    blo

    od

    lea

    ves

    the

    con

    us

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    rio

    sus,

    it

    mo

    ves

    thro

    ug

    h

    the

    gil

    ls,

    bec

    om

    ing

    oxy

    gen

    ated

    . B

    loo

    d l

    eavi

    ng

    th

    e g

    ills

    th

    en

    flo

    ws

    thro

    ug

    h a

    net

    wo

    rk o

    f ar

    teri

    es t

    o t

    he

    rest

    of

    the

    bo

    dy,

    fi-

    nal

    ly r

    etu

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    g t

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    he

    sin

    us

    ven

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    his

    sim

    ple

    lo

    op

    has

    on

    e se

    rio

    us

    lim

    itat

    ion

    : in

    pas

    sin

    g t

    hro

    ug

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    he

    cap

    illa

    ries

    in

    th

    e g

    ills

    , blo

    od

    pre

    ssu

    re d

    rop

    s si

    gn

    ific

    antl

    y. T

    his

    slo

    ws

    circ

    ula

    tio

    n

    fro

    m t

    he

    gil

    ls t

    o t

    he

    rest

    of

    the

    bo

    dy

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    can

    lim

    it o

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    en d

    e-li

    very

    to

    tis

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    cha

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    r 5

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    a.

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    Y # W

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    In a

    mp

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    ian

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    nd

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    ep

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    s, lu

    ng

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    req

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    Th

    e ad

    ven

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    am

    ph

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    ch

    apte

    r 4

    9)

    invo

    lved

    a

    maj

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    ge

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    he

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    tern

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    er b

    loo

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    s p

    um

    ped

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    he

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    ot

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    ectl

    y to

    th

    e ti

    ssu

    es o

    f th

    e b

    od

    y. I

    nst

    ead

    , it

    is

    retu

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    th

    e pu

    lmon

    ary

    vein

    s to

    th

    e h

    eart

    . B

    loo

    d l

    eave

    s th

    e h

    eart

    a s

    eco

    nd

    tim

    e to

    be

    circ

    ula

    ted

    th

    rou

    gh

    oth

    er ti

    ssu

    es. T

    his

    syst

    em is

    term

    ed d

    ou

    ble

    cir

    cula

    tio

    n:

    On

    e sy

    stem

    , th

    e p

    ulm

    on

    ary

    circ

    ula

    tio

    n, m

    ove

    s b

    loo

    d b

    etw

    een

    h

    eart

    an

    d l

    un

    gs,

    an

    d a

    no

    ther

    , th

    e sy

    stem

    ic c

    ircu

    lati

    on

    , m

    ove

    s b

    loo

    d b

    etw

    een

    th

    e h

    eart

    an

    d t

    he

    rest

    of

    the

    bo

    dy.

    Am

    ph

    ibia

    n c

    ircu

    lati

    on

    Op

    tim

    ally

    , o

    xyg

    enat

    ed b

    loo

    d f

    rom

    lu

    ng

    s w

    ou

    ld g

    o d

    irec

    tly

    to

    tiss

    ues

    , ra

    ther

    th

    an b

    ein

    g m

    ixed

    in

    th

    e h

    eart

    wit

    h d

    eoxy

    gen

    -at

    ed b

    loo

    d r

    etu

    rnin

    g f

    rom

    th

    e b

    od

    y. T

    he

    amp

    hib

    ian

    hea

    rt h

    as

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    str

    uct

    ura

    l fe

    atu

    res

    that

    sig

    nif

    ican

    tly

    red

    uce

    th

    is m

    ixin

    g

    (fig

    ure

    50

    .6) .

    Fir

    st,

    the

    atri

    um

    is

    div

    ided

    in

    to t

    wo

    ch

    amb

    ers:

    T

    he

    rig

    ht

    atri

    um

    rec

    eive

    s d

    eoxy

    gen

    ated

    blo

    od

    fro

    m t

    he

    sys-

    tem

    ic

    circ

    ula

    tio

    n,

    and

    th

    e le

    ft

    atri

    um

    re

    ceiv

    es

    oxy

    gen

    ated

    b

    loo

    d f

    rom

    th

    e lu

    ng

    s. T

    hes

    e tw

    o t

    ypes

    of

    blo

    od

    , th

    eref

    ore

    , do

    n

    ot

    mix

    in

    th

    e at

    ria.

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    as a

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    tric

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    y an

    d s

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    mic

    cir

    cula

    tio

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    is i

    n-

    com

    ple

    te.

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    e ex

    ten

    t o

    f m

    ixin

    g w

    hen

    th

    e co

    nte

    nts

    of

    each

    at

    riu

    m e

    nte

    r th

    e ve

    ntr

    icle

    is

    red

    uce

    d b

    y in

    tern

    al c

    han

    nel

    s cr

    e-at

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    y re

    cess

    es i

    n t

    he

    ven

    tric

    ula

    r w

    all.

    Th

    e co

    nu

    s ar

    teri

    osu

    s is

    p

    arti

    ally

    sep

    arat

    ed b

    y a

    div

    idin

    g w

    all,

    wh

    ich

    dir

    ects

    deo

    xyg

    en-

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    blo

    od

    in

    to t

    he

    pu

    lmo

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    y ar

    teri

    es a

    nd

    oxy

    gen

    ated

    blo

    od

    in

    to t

    he

    aort

    a, t

    he

    maj

    or

    arte

    ry o

    f th

    e sy

    stem

    ic c

    ircu

    lati

    on

    .

    Am

    ph

    ibia

    ns

    livi

    ng

    in

    wat

    er c

    an o

    bta

    in a

    dd

    itio

    nal

    oxy

    gen

    b

    y d

    iffu

    sio

    n t

    hro

    ug

    h t

    hei

    r sk

    in. T

    hu

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    hib

    ian

    s h

    ave

    a pu

    lmo-

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    uit

    th

    at s

    end

    s b

    loo

    d t

    o b

    oth

    th

    e lu

    ng

    s an

    d t

    he

    skin

    . C

    uta

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    us

    resp

    irat

    ion

    is a

    lso

    see

    n in

    man

    y aq

    uat

    ic r

    epti

    les

    such

    as

    tu

    rtle

    s.

    Re

    pti

    lia

    n c

    ircu

    lati

    on

    Am

    on

    g r

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    dit

    ion

    al m

    od

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    atio

    ns

    hav

    e fu

    rth

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    ced

    th

    e m

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    g o

    f b

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    th

    e h

    eart

    . In

    ad

    dit

    ion

    to

    hav

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    tw

    o s

    ep-

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    e at

    ria,

    rep

    tile

    s h

    ave

    a se

    ptu

    m t

    hat

    par

    tial

    ly s

    ub

    div

    ides

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    e ve

    ntr

    icle

    . Th

    is s

    epar

    atio

    n i

    s co

    mp

    lete

    in

    on

    e o

    rder

    of

    rep

    tile

    s,

    the

    cro

    cod

    ilia

    ns,

    wh

    ich

    hav

    e tw

    o s

    epar

    ate

    ven

    tric

    les

    div

    ided

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    a co

    mp

    lete

    sep

    tum

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    e th

    e fo

    llo

    win

    g s

    ecti

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    ). A

    no

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    e in

    th

    e ci

    rcu

    lati

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    of

    rep

    tile

    s is

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    at t

    he

    con

    us

    arte

    rio

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    has

    b

    eco

    me

    inco

    rpo

    rate

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    nto

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    e tr

    un

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    f th

    e la

    rge

    arte

    ries

    lea

    v-in

    g t

    he

    hea

    rt.

    Fig

    ure

    50

    .6 T

    he

    he

    art

    an

    d c

    ircu

    lati

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    of

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    am

    ph

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    d b

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    lin

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    spec

    tive

    ly)

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    lit

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    as t

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    mp

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    o t

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    lu

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    s. O

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    enat

    ion

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    od

    als

    o o

    ccu

    rs b

    y g

    as e

    xch

    ang

    e

    thro

    ug

    h t

    he

    skin

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    10

    24

    p

    art

    VII

    An

    ima

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    rm a

    nd

    Fu

    nct

    ion

    Apago PDF Enhancer

    Head

    Body

    Lungs

    Systemic

    capillaries

    Pulm

    onary

    artery

    Pulm

    onary

    semilu

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    lve

    Inferior

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    Superior

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    Tricuspid valve

    Right ve

    ntricle

    Left

    ventricle

    Right atrium

    Pulm

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    veins

    Left atrium

    Bicuspid

    mitral

    valve

    Systemic

    capillaries

    Pulm

    onary

    artery

    Superior

    vena cava

    Inferior

    vena cava

    Artery

    Pulm

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    Aorta

    Respiratory

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    T

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    Alt

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    ev

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    .3

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    Fig

    ure

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    fig

    ure

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    and

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    10

    26

    p

    art

    VII

    An

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    l Fo

    rm a

    nd

    Fu

    nct

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    Apago PDF Enhancer

  • Apago PDF Enhancer

    F ig

    ure

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    s

    10

    28

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    VII

    An

    ima

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    rm a

    nd

    Fu

    nct

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    Apago PDF Enhancer

    Fig

    ure

    50

    .11

    Me

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    pre

    ssu

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    s o

    f th

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    , eve

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    imu

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    nd

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    as

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    T

    he

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    d a

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    ran

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    50

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    vein

    s em

    pty

    in

    to t

    he

    rig

    ht

    atri

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    , co

    m-

    ple

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    he

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    n.

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    Do

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    p t

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    a.

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