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C
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1
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1
I
n
tr
o
d
u
c
ti
o
n
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3
1.
2
A
n
at
o
m
y
of
th
e
H
e
ar
t
..
..
..
..
..
..
..
..
..
3
1.
2.
1
C
h
a
m
b
er
a
n
d
V
al
v
es
..
..
..
..
..
..
..
.
.
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.
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4
1
.
2
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2
C
a
r
d
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a
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C
e
ll
a
n
d
C
a
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a
c
M
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4
1
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2
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3
C
o
ro
n
ar
y
A
rt
er
ie
s
a
n
d
C
ar
di
a
c
V
ei
n
s
..
..
.
6
1.
2.
4
V
e
n
o
u
s
C
ir
c
ul
at
io
n
..
..
..
..
..
..
..
..
..
..
.
6
1.
2.
5
N
e
r
v
e
S
u
p
p
l
y
o
f
t
h
e
H
e
a
rt
.
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9
1
.
2
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6
C
1.3.1
o
n
d
u
c
ti
o
n
S
y
st
e
m
of
th
e
H
e
ar
t
..
..
..
..
.
1
0
1.
3
P
h
y
si
ol
o
g
y
of
th
e
H
e
ar
t
..
..
..
..
..
..
..
..
1
1
C
ir
c
ul
at
or
y
S
y
st
e
m
:
S
y
st
e
m
i
c
a
n
d
P
u
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m
o
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a
r
y
C
ir
c
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l
a
ti
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n
.
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1
1
1
.
3
.
2
C
o
n
d
u
ct
io
n
S
y
st
e
m
of
th
e
H
e
ar
t
(
E
x
ci
ta
ti
o
n
S
e
q
u
e
n
c
e)
..
..
..
..
..
..
..
..
..
1
1
1.
3.
3
A
ct
io
n
P
ot
e
nt
ia
l
(
A
P
)
.
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1
1
1.3.4
M
e
c
h
a
n
is
m
o
f
E
x
c
it
a
ti
o
n
a
n
d
C
o
n
tr
a
c
ti
o
n
C
o
u
pl
in
g
of
C
ar
di
a
c
M
y
o
c
yt
es
..
..
..
..
..
.
1
3
1.
3.
5
A
ut
o
n
o
m
ic
N
er
v
o
u
s
S
y
st
e
m
a
n
d
H
e
ar
t
.
.
.
.
1
6
1
.
3
.
6
C
a
r
d
i
a
c
C
y
c
l
e
.
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1
6
1
.
3
.
7
P
h
y
si
ol
o
g
y
of
C
or
o
n
ar
y
C
ir
c
ul
at
io
n
..
..
..
1
7
1.
3.
8
C
or
o
n
ar
y
C
ol
la
te
ra
ls
..
..
..
..
..
..
..
..
..
.
1
7
1
.
4
E
x
e
r
c
is
e
P
h
y
si
o
l
o
g
y
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1
8
1
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4
.
1
G
e
n
d
e
r
a
n
d
E
x
er
ci
se
P
er
fo
r
m
a
n
c
e
..
..
..
.
1
9
1.
4.
2
A
g
e
a
n
d
E
x
er
ci
se
P
er
fo
r
m
a
n
c
e
..
..
..
..
..
1
9
1.
5
C
o
10
n
c
l
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i
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2
0
R
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f
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s
.
.
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.
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.
.
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.
.
..
..
..
..
..
..
..
.
2
0
1
.
1
I
n
t
r
o
d
u
c
t
i
o
n
T
h
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i
m
p
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o
f
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w
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c
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A
n
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V
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d
u
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i
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g
t
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1
6
t
h
c
e
n
t
u
r
y
[
1
]
a
n
d
f
r
o
m
b
i
r
t
h
t
o
d
e
a
t
h
,
t
h
e
h
e
a
r
12
i
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t
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t
h
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m
o
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t
t
a
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k
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d
a
b
o
u
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f
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h
u
m
a
n
b
o
d
y
.
I
t
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c
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n
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r
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m
a
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y
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t
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l
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s
,
s
u
c
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a
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p
h
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p
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s
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a
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s
,
p
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s
a
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d
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s
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o
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T
h
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m
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a
n
14
s
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b
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t
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16
i
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.
18
1
.
2
A
n
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H
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T
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a
n
20
d
w
e
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g
h
t
(
T
a
b
l
e
1
.
1
)
.
T
h
e
t
i
p
(
a
p
e
x
)
o
f
n
t
e
d
f
o
r
w
a
r
d
,
d
o
w
n
w
a
r
d
,
a
n
d
t
o
w
a
r
d
t
h
e
t
h
e
l
e
f
t
.
h
e
a
r
t
T
h
e
i
s
p
o
i
(
i
n
f
e
r
i
o
r
)
d
i
a
p
h
r
a
g
m
a
t
i
c
s
u
r
f
a
c
e
l
i
e
s
d
i
r
e
c
t
l
y
o
n
t
h
e
d
i
a
p
h
r
a
g
m
.
T
h
e
h
e
a
r
t
l
i
e
s
i
n
a
d
o
u
b
l
e
w
a
l
l
e
d
f
i
b
r
o
s
e
r
o
u
s
s
a
c
c
a
22
l
l
e
d
t
h
e
p
e
r
i
c
a
r
d
i
a
l
s
a
c
,
w
h
i
c
h
i
s
d
i
v
i
d
e
d
i
n
t
o
(
a
)
f
i
b
r
o
u
s
p
e
r
i
c
a
r
d
i
u
m
,
a
n
d
(
b
)
s
e
r
o
u
s
p
e
r
i
c
a
r
d
i
u
m
.
T
h
e
f
i
b
r
o
u
s
p
e
r
i
c
a
r
d
i
u
m
r
e
a
t
v
e
s
s
e
l
s
e
n
v
e
l
o
p
s
[
2
]
.
t
h
e
s
e
r
o
u
s
h
e
a
r
t
a
n
d
a
t
t
a
c
h
e
s
T
h
e
p
e
r
i
c
a
r
d
i
u
m
i
s
a
o
n
t
o
t
h
e
g
c
l
o
s
e
d
s
a
24
c
c
o
n
s
i
s
t
i
n
g
o
f
t
w
o
l
a
y
e
r
s
a
v
i
s
c
e
r
a
l
l
a
y
e
r
o
r
e
p
i
c
a
r
d
i
u
m
f
o
r
m
i
n
g
t
h
e
o
u
t
e
r
l
i
n
i
n
g
o
f
t
h
e
g
r
e
a
t
v
e
s
s
e
l
s
a
n
d
t
h
h
e
a
r
t
,
t
h
e
a
n
d
f
i
b
r
o
u
s
a
p
a
r
i
e
t
a
l
l
a
y
e
r
f
o
r
m
i
n
g
a
n
i
n
n
e
r
l
i
n
i
n
g
o
p
e
r
i
c
a
r
d
i
u
m
[
2
4
]
.
T
h
e
t
w
o
l
a
y
e
r
s
o
f
t
h
e
26
s
e
r
o
u
s
p
e
r
i
c
a
r
d
i
u
m
c
o
n
t
a
i
n
t
h
e
p
e
r
i
c
a
r
d
i
a
l
f
l
u
i
d
,
w
h
i
c
h
p
r
e
v
e
n
t
s
f
r
i
c
t
i
o
n
b
e
t
w
e
e
n
t
h
e
h
e
a
r
t
a
n
d
t
h
e
p
e
r
i
c
a
r
d
i
u
m
[
2
4
]
.
Ta
ble
1.1
An
ato
mi
cal
fac
ts
ab
out
the
hu
ma
n
he
art
Nor
mal
hum
an
hear
t
vari
es
with
heig
ht
and
wei
ght
Wei
ghs
appr
oxi
mat
ely
300
350
gra
ms
in
mal
es
Wei
ghs
appr
oxi
mat
ely
250
300
gra
ms
in
fem
ales
Rig
ht
ven
tricl
e
thic
kne
ss is
0.3
0.5
cm
Left
ven
tricl
e
thic
kne
ss is
1.3
1.5
cm
Div
ide
d
into
four
dist
inct
cha
mb
ers
Co
mp
ose
d of
thre
e
laye
rs
(epi
car
diu
m,
28
myo
card
ium
and
end
ocar
diu
m)
Con
tain
s
two
atria
(left
and
righ
t)
Con
tain
s
two
vent
ricle
s
(left
and
righ
t)
Con
tain
s
four
valv
es
(aor
tic,
mitr
al,
tricu
spid
,
pul
mon
ary)
T
he
wa
ll
of
th
e
he
art
is
co
m
po
se
d
of
th
re
e
la
ye
rs:
(a
)
ep
ic
ar
di
u
m;
(b
)
m
yo
ca
rd
iu
m;
an
d
(c
)
en
do
ca
rd
iu
m
(F
ig.
1.
1)
[5
,
6]
.
T
he
ep
ic
ar
di
u
m
is
th
e
ou
ter
lin
in
g
of
th
e
ca
rdi
ac
ch
a
m
be
rs
an
d
is
for
m
ed
by
th
e
vis
ce
ral
la
ye
r
of
th
e
ser
ou
s
pe
ric
ar
di
u
m.
Th
e
m
yo
ca
rdi
u
m
is
th
e
int
er
m
ed
iat
e
la
ye
r
of
th
e
he
art
an
d
is
co
m
po
se
d
of
th
re
e
di
sc
er
na
bl
e
la
ye
rs
of
m
us
cl
e
[5
,
6]
th
at
ar
e
se
en
pr
ed
o
mi
30
na
ntl
y
in
th
e
lef
t
ve
ntr
icl
e
an
d
int
erve
ntr
ic
ul
ar
se
pt
u
m
al
on
e
an
d
in
cl
ud
es
a
su
be
pi
ca
rdi
al
la
ye
r,
a
mi
dd
le
co
nc
en
tri
c
la
ye
r
an
d
a
su
be
nd
oc
ar
di
al
la
ye
r.
T
he
re
st
of
th
e
he
art
is
co
m
po
se
d
m
ai
nl
y
of
th
e
su
be
pi
ca
rd
ial
an
d
su
be
nd
oc
ar
di
al
la
ye
rs
[7
,
8]
.
Th
e
m
yo
ca
rdi
u
m
als
o
co
nt
ai
ns
im
po
rta
nt
str
uc
tur
es
su
ch
as
ex
cit
ab
le
no
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l
tis
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th
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tin
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ste
m.
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e
en
do
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rdi
u
m
th
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r
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os
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la
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r
of
th
e
he
art
is
fo
r
m
ed
of
th
e
en
do
th
eli
u
m
an
d
su
be
nd
ot
he
lia
l
co
nn
ec
tiv
e
tis
su
e
[5
,
6]
.
32
a
r
t
i
s
d
i
Endocardium
v
i
d
eMyocar
d
i
n
Fi
t
g
o
1.1
f
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o
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of
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i
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n
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2
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h
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t
t
h
i
c
k
n
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s
s
[
2
,
4
,
9
]
.
T
h
e
l
e
f
t
a
t
r
i
u
m
(
L
A
)
a
n
d
r
i
g
h
t
a
t
r
i
u
m
(
R
A
)
a
r
e
s
m
a
l
l,
t
h
i
n
w
a
l
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d
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a
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s
l
o
c
a
t
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d
j
u
s
t
a
b
o
v
e
t
h
e
l
34
e
f
t
v
e
n
t
r
i
c
l
e
(
L
V
)
a
n
d
r
i
g
h
t
v
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n
t
r
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c
l
e
(
R
V
)
,
r
e
s
p
e
c
t
i
v
e
l
y
.
T
h
e
v
e
n
t
r
i
c
l
e
s
a
r
e
l
a
r
g
e
r
t
h
i
c
k
w
a
l
l
e
d
c
h
a
m
b
e
r
s
t
h
a
t
p
e
r
f
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r
m
m
o
s
t
o
f
t
h
e
w
o
r
k
[
2
,
4
,
9
]
(
T
a
b
l
e
1
.
2
)
.
T
h
e
a
t
r
i
a
r
e
c
e
i
v
e
b
l
o
o
d
f
r
o
m
t
h
e
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o
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s
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y
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t
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m
a
n
d
l
u
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g
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d
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h
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n
c
o
n
t
r
a
c
t
a
n
d
e
j
e
c
t
t
h
e
b
l
36
o
o
d
i
n
t
o
t
h
e
v
e
n
t
r
i
c
l
e
s
.
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h
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t
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c
l
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t
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n
p
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m
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l
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o
d
t
h
r
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g
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t
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y
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l
u
n
g
s
.
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h
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h
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a
r
t
c
o
n
t
a
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f
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r
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l
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n
d
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u
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k
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n
o
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t
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m
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b
r
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p
t
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m
,
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o
r
t
i
c
i
n
t
e
r
v
a
l
v
u
l
a
r,
r
i
38
g
h
t
,
a
n
d
l
e
f
t
f
i
b
r
o
u
s
t
r
i
g
o
n
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s
[
3
,
4
,
6
,
1
0
,
1
1
]
(
F
i
g
.
1
.
2
,
T
a
b
l
e
1
.
3
)
.
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h
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t
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r
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d
t
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m
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b
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a
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o
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s
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t
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f
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t
r
a
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f
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b
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u
s
b
o
d
y
,
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h
i
c
h
i
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p
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n
e
t
r
a
t
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d
b
y
t
h
e
b
u
n
d
l
e
o
f
H
i
s
[
3
,
4
,
6
,
1
0
,
1
1
]
.
T
h
e
f
i
b
r
o
u
s
s
k
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l
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t
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n
f
u
n
c
t
i
o
n
s
n
o
t
o
n
l
y
t
o
p
r
o
40
v
i
d
e
a
n
e
l
e
c
t
r
o
p
h
y
s
i
o
l
o
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t
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p
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t
[
8
,
1
2
,
1
3
]
.
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a
c
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t
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c
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s
t
a
b
i
l
i
t
y
[
3
]
.
42
1
.
2
.
2
C
a
r
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(
d
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l
a
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d
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r
m
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n
a
l
s
)
[
6
,
1
0
,
1
1
,
1
4
1
6
]
.
T
h
e
s
a
r
c
44
o
m
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s
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t
h
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c
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o
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l
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s
,
w
h
i
c
h
c
o
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d
u
c
t
t
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c
t
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n
p
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e
n
t
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a
l
(
A
P
)
t
o
t
h
e
i
n
t
e
r
i
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o
f
t
h
e
c
e
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l
[
6
,
1
4
]
.
T
h
e
T
t
u
b
u
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s
c
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n
e
c
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a
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c
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m
m
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t
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t
h
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s
a
r
c
46
o
p
l
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s
m
i
c
r
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t
i
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u
m
i
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t
h
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k
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d
t
h
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c
a
r
d
i
a
c
m
u
s
c
l
e
[
1
4
,
1
5
]
.
C
a
r
d
i
a
c
m
u
s
c
l
e
i
s
a
n
i
n
v
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l
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d
m
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,
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a
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r
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a
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f
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m
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t
e
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n
f
il
a
m
e
n
t
s
,
w
h
i
c
h
a
r
e
48
a
n
c
h
o
r
e
d
b
y
s
e
g
m
e
n
t
s
c
a
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d
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l
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s
.
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a
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a
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m
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s
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a
t
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o
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t
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k
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t
a
l
m
u
s
c
l
e
[
6
,
1
0
,
1
1
,
1
4
1
6
]
a
n
d
a
c
ti
n
a
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d
m
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.
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h
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50
p
p
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a
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a
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g
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b
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d
s
,
w
h
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e
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b
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n
d
s
[
8
,
1
2
,
1
3
,
1
5
].
T
h
e
d
a
r
k
b
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d
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,
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,
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3
,
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5
].
T
h
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t
h
i
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n
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r
52
a
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t
r
a
c
ti
o
n
[
6
,
1
4
,
1
5
]
.
C
a
r
d
i
a
c
m
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s
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b
a
n
d
s
(
s
p
e
c
i
a
l
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s
e
d
54
56
T
Left ventricle (LV)
1. Made of an inlet portion comprised of mitral valve apparatus, subaortic outflow portion and a trabeculated apical zone
2. Three times thicker than the RV and most muscular
3. Thickest towards the base and thinnest towards the apex
4. LV free wall and septal thickness is three times the thickness of the RV free wall
5. Mitral and aortic valves share fibrous continuity
6. LV apex is relatively less trabeculated than the RV apex
Right ventricle (RV)
1. Comprised of inlet and outflow segments
2. Inlet extends from tricuspid annulus to the insertions of the papillary muscles
3. Apical trabecular zone extends inferiorly beyond the papillary muscle attachment toward the ventricular apex and halfway
along the anterior wall.
4. Outflow portion (conus) is a muscular subpulmonary channel
5. Arch shaped muscular ridge separates the tricuspid and pulmonary valves
Right atrium (RA)
1. Thinnest walls of the four chambers
2. Forms the right border of the heart
3. Gives off the right auricular appendage
4. Receives the superior vena cava, inferior vena cava and coronary sinus
5. Discharges into right ventricle through the tricuspid valve
Left atrium (LA)
1. Forms base of the heart (posterior surface)
2. Gives off the left auricular appendage.
3. Receives two right pulmonary veins (sometimes three) and two left pulmonary veins (sometimes one)
4. Discharges into the left ventricle through the mitral valve
Ventricular septum
Intracardiac partition having four parts (inlet, membranous, trabecular and infundibular)
Divided into muscular and membranous septum
Membranous septum lies beneath the right and posterior aortic cusps and contact mitral and tricuspid annuliAtrial septum
b
l
e
1
.
2
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a
r
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a
c
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a
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c
ul
ar
c
h
a
m
b
er
s
[
2,
4
6,
9
]
m
b
e
Righ
r
s
Tricuspid
a
n
Right ve
d
F
i
g
1
.
2
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a
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j
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c
t
i
o
n
s
58
)
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h
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s
[
6
,
1
4
,
1
5
]
a
n
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i
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60
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1
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,
[
6
,
1
5
]
.
62
h
i
s
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f
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t
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a
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h
e
[
8
,
i
n
d
i
v
i
d
u
a
l
1
2
1
5
]
.
c
e
l
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1
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C
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V
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C
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1
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]
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F
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.
1
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[
2
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2
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h
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[
2
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70
C
x
3
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a
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s
8
]
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w
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R
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[
2
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3
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A
V
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[
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]
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(
8
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%
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(
A
V
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)
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i
n
a
l
l
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,
t
h
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c
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n
a
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a
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s
b
r
a
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c
76
h
i
n
t
o
s
m
a
l
l
i
n
a
t
e
i
n
e
n
d
a
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d
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h
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r
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o
l
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s
.
T
h
e
s
e
v
e
s
s
e
l
s
t
e
r
m
s
u
p
p
l
y
t
h
e
m
y
o
c
a
r
d
i
a
l
t
i
s
s
u
e
w
i
t
h
b
l
o
o
d
[
2
,
3
,
6
8
]
.
I
n
ge
ner
al,
the
RC
A
is
do
mi
na
nt
in
60
65
%
of
cas
es
bec
aus
e it
giv
es
off
a
PD
A
bra
nc
h
(ba
lan
ced
cor
on
ary
cir
cul
ati
on)
[2,
3,
6
8].
In
ab
out
10
15
%
of
cas
es,
the
LC
x
giv
es
ris
e
to
the
PD
A
(lef
t
pre
do
mi
na
nt
cir
cul
ati
on)
. In
20
25
%
of
cas
es,
the
78
RC
A,
in
ad
diti
on
to
sup
ply
ing
the
PD
A,
cro
sse
s
the
pos
teri
or
int
erv
ent
ric
ula
r
sep
tu
m
to
rea
ch
as
far
as
the
left
ma
rgi
nal
art
ery
an
d
the
reb
y
sup
ply
the
dia
phr
ag
ma
tic
sur
fac
e
of
the
left
ve
ntri
cle
(ri
ght
pre
do
mi
na
nce
)
[2,
3,
8].
Ho
we
ver
,
thi
s
ter
m
do
es
not
dis
tin
gui
sh
thi
s
co
ndi
tio
n
fro
m
bal
anc
ed
cor
on
ary
cir
cul
ati
on
(Ta
ble
1.5
)
[7]
.
1
.
2
.
4
V
e
n
o
u
s
C
i
r
c
u
l
a
t
i
o
n
Th
e
ve
no
us
cir
cul
ati
on
of
the
he
art
is
fro
m
the
cor
on
ary
sin
us,
ant
eri
or
car
dia
c
vei
ns
an
d
the
les
ser
car
dia
c
(th
eb
esi
an)
vei
ns
[6]
.
Th
e
cor
on
ary
sin
us
rec
eiv
es
mo
st
of
the
ve
no
us
ret
urn
fro
m
the
epi
car
diu
m
an
d
my
oc
ard
iu
m
[6]
an
80
d it
op
ens
int
o
the
rig
ht
atri
um
bet
we
en
the
op
eni
ng
of
the
inf
eri
or
ve
na
ca
va
an
d
the
rig
ht
AV
val
ve
[2,
3,
5,
6].
Th
e
cor
on
ary
sin
us
giv
es
ris
e
to
tri
but
ari
es
suc
h
as
the
gre
at
car
dia
c
vei
n,
mi
ddl
e
car
dia
c
vei
n,
sm
all
er
car
dia
c
vei
n
an
d
obl
iqu
e
vei
n.
Th
e
gre
at
car
dia
c
vei
n
dra
ins
the
ant
eri
or
por
tio
n
of
the
int
erv
ent
ric
ula
r
sep
tu
m
an
d
ant
eri
or
asp
ect
s
of
bot
h
ve
ntr
icl
es
[2
6].
T
he
mi
ddl
e
car
dia
c
vei
n
dra
ins
the
pos
teri
or
por
tio
n
of
the
int
erv
ent
ric
ula
r
sep
tu
m
an
d
pos
teri
or
asp
ect
of
SA n
Right c
artery
AV n
Bu
of
R
v
Fig
1.3
Cor
ona
ry
arte
ries
82
Tab
Tricuspid valve
1. Tricuspid valve (right atrioventricular valve) connects the right atrium and the right ventricle
2. Composed of five components (annulus, leaflets, commissures, chordae tendineae, papillary muscles)
3. Anterior tricuspid leaflet is the largest (most mobile) and the posterior leaflet is the smallest
4. Tricuspid valve has a triangular orifice
Mitral valve
Mitral valve (left atrioventricular valve) connects the left atrium to the left ventricle
Composed of five components (annulus, leaflets, commissures, chordae tendineae, papillary muscles) 3. Composed
of two leaflets only
4. Anterior tricuspid leaflet is large, semicircular and twice the height of posterior leaflet
5. Posterior leaflet is rectangular and is divided into three scallops
6. Mitral valve has an elliptical orifice
Aortic valve
1. Aortic valve (semilunar valve) opens between the left ventricle and the aorta
2. Composed of three components (annulus, cusps, commissures)
3. Composed of three semilunar cusps
Pulmonary valves
1. Pulmonary valve (semilunar valve) regulates flow between the right ventricle and the pulmonary artery
2. Composed of three components (annulus, cusps, commissures)
Cardiac grooves
1. Atrioventricular (AV) groove separates the atria from the ventricle
2. Anterior and posterior interventricular (IV) grooves separate the ventricles
3. Right coronary arteries (RCA) travel in the right atrioventricular groove
4. Circumflex artery (Cx) travels in the left atrioventricular groove
5. The left anterior descending artery (LAD) travels along the anterior interventricular groove
6. Posterior descending artery (PDA) travels along posterior interventricular groove
Cardiac crux (external and internal)
1. External cardiac crux is the intersection between the AV, posterior IV and interatrial (IA) grooves
2. Internal cardiac crux is the posterior intersection between the mitral and tricuspid annuli and the atrial and ventricular
septa
Cardiac margins
1. Acute margin junction between anterior and inferior wall of the right ventricle
2. Obtuse margin rounded lateral wall of left ventricle
84
1.3 c
Car skel
dia eto
c
n
val [2,
ves 4
and 6,
car 8,
dia 9]
Table
1.4
Coronary
blood supply
[2, 3, 6, 8]
Coronary arterial ci
B. Right coronary ar
1. Arises from the right
C. Left circumflex a
Coronary collaterals
1. Provide communicati
Table
1.4
(continued)
Coronary blood
supply [2, 3, 6, 8]
Great vessels
Right ventricle
Right atrium
Diaphragmatic surface/i
left ventricle (LV)
bot
h
ve
ntr
icl
es
[4,
6]
an
d
the
sm
all
er
car
dia
c
vei
n
dra
ins
the
ma
r
gin
al
asp
ect
of
the
rig
ht
ve
ntr
icl
e
[11
].
Th
e
the
bes
ian
vei
ns
dra
in
the
en
do
car
diu
m
an
d
the
inn
er
mo
st
lay
ers
of
the
my
oc
ard
iu
m
dir
ect
ly
int
o
the
un
der
lyi
ng
ch
am
ber
[11
].
1
.
2
.
5
N
e
r
v
e
S
u
p
p
l
y
o
f
t
h
e
H
e
a
r
t
Th
e
sy
mp
ath
eti
c
an
d
par
asy
mp
ath
eti
c
aut
on
om
ic
ner
vo
us
su
ppl
ies
to
the
he
art
for
m
the
car
dia
c
ple
xu
s,
wh
ich
is
loc
ate
d
clo
se
to
the
arc
h
of
the
aor
ta.
Th
e
fib
res
fro
m
the
car
dia
c
ple
xu
s
acc
om
pa
ny
the
cor
on
ary
art
eri
es
an
d
rea
ch
the
he
art,
wit
h
mo
st
of
the
m
ter
mi
nat
ing
at
the
SA
no
de,
AV
no
de
an
d a
mu
ch
les
s
de
nse
su
ppl
y
to
the
atri
al
an
d
ve
ntr
icu
lar
my
oc
ard
iu
m
[11
].
In
ge
ner
al,
the
par
asy
mp
ath
eti
c
va
gal
fib
res
are
inh
ibit
ory
an
d
red
uc
e
the
he
art
rat
e
an
d
str
ok
e
vol
um
e.
Th
e
sy
mp
ath
eti
c
ner
ves
act
as
acc
ela
tor
y
ner
ves
inc
rea
sin
g
bot
h
the
he
art
rat
e
an
d
str
ok
e
vol
um
e
[11
].
Th
e
aff
ere
nt
ner
ves
run
alo
ng
sy
mp
ath
eti
c
pat
hw
ays
via
bot
h
car
dia
c
acc
ele
rat
or
ner
ves
an
d
tho
rac
ic
spl
an
ch
nic
ner
ves
to
rea
ch
the
int
er
me
dio
lat
era
l
hor
n
of
T1
T4
of
the
spi
nal
cor
d
[11
].
Th
e
nor
adr
en
erg
ic
or
the
sy
mp
ath
eti
c
ner
vo
us
sys
te
m
is
ma
inl
y
inv
olv
ed
wit
h
inc
rea
sin
g
the
he
art
rat
e
(ch
ron
otr
op
y),
co
ntr
act
ilit
y
(io
not
rop
y)
an
d
the
spe
ed
of
co
nd
uct
ion
(dr
om
otr
op
y)
in
the
car
dia
c
mu
scl
e
fib
res
an
d
the
co
nd
uct
ion
tiss
ue;
an
d
the
tra
ns
mit
ter
inv
olv
ed
is
ma
inl
y
nor
epi
ne
phr
ine
[6]
.
Th
e
SA
no
de
rec
eiv
es
mo
st
of
it
ner
ve
fib
res
fro
m
the
rig
htsid
ed
tho
rac
ic
sy
mp
ath
eti
c
ga
ngl
ia
an
d
the
rig
ht
va
gu
s
[8]
.
Th
e
AV
no
des
an
d
ve
ntr
icl
es
rec
eiv
e
the
ir
ner
ve
su
ppl
y
for
m
the
left
sid
ed
tho
rac
ic
sy
mp
ath
eti
c
ga
ngl
ia
an
d
the
left
11
va
gu
s,
wh
ich
is
ma
inl
y
be
ca
use
SA
no
de
de
vel
op
s
fro
m
the
str
uct
ure
s
on
the
rig
ht
sid
e
of
the
em
bry
o
an
d
the
AV
no
de
de
vel
op
s
fro
m
the
str
uct
ure
s
on
the
left
sid
e
of
the
em
bry
o
[8]
.
T
h
e
s
y
m
p
a
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h
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t
i
c
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f
f
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t
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h
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c
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p
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r
s
,
w
h
i
c
h
i
n
c
l
u
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e
s
1
a
n
d
2
a
d
r
e
n
e
r
g
i
c
r
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c
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p
t
o
r
s
[
2
,
1
1
]
.
1
r
e
c
e
p
t
o
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r
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f
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u
n
d
m
a
13
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n
l
y
i
n
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S
A
n
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d
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V
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A
M
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(
c
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)
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d
s
y
m
15
p
a
t
h
e
t
i
c
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f
f
e
c
t
s
[
2
,
1
1
]
.
2
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p
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.
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a
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o
f
1
r
e
17
c
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p
t
o
r
s
,
l
a
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a
t
i
o
n
i
.
e
.
,
o
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g
u
l
a
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o
f
b
l
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d
f
l
o
w
[
2
,
1
1
]
.
T
h
e
c
h
o
l
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i
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p
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c
b
l
o
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d
p
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n
e
r
v
o
u
s
19
s
y
s
t
e
m
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f
f
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t
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a
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d
a
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a
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d
t
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r
a
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m
i
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v
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l
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d
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m
a
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n
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t
y
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c
h
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l
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t
i
c
f
i
b
r
e
s
[
8
,
t
o
1
2
,
t
h
e
1
3
]
.
h
e
a
r
t
T
h
e
v
a
g
i
s
u
p
p
l
y
t
h
e
p
a
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p
a
t
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a
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c
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c
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l
e
x
u
s
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s
.
T
h
e
21
p
a
r
a
s
y
m
p
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f
f
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t
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d
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a
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a
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m
u
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c
a
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c
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p
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o
r
s
,
w
h
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a
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b
y
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A
M
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,
c
o
n
t
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a
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d
c
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d
u
23
c
t
i
o
n
v
e
l
o
c
i
t
y
[
8
,
1
2
,
1
3
]
.
T
h
e
a
u
t
o
n
o
m
i
c
c
e
n
t
r
e
s
C
N
S
,
m
a
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n
l
y
t
h
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v
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s
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m
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m
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a
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y
p
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c
a
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d
i
o
v
a
25
s
c
u
l
a
r
s
y
s
t
e
m
,
d
e
p
e
n
d
i
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o
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a
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t
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p
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r
i
p
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y
a
n
d
t
h
e
C
N
S
[
7
,
8
,
1
2
,
1
3
]
.
T
h
e
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s
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m
a
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v
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s
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m
p
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t
i
c
n
e
27
r
v
e
s
[
7
,
8
,
1
2
,
1
3
]
.
B
o
t
h
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h
e
s
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m
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C
N
S
29
,
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[
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,
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.
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m
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31
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6
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33
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.
T
h
e
s
e
a
r
e
i
m
p
o
r
t
a
n
t
i
n
m
a
i
n
t
a
i
n
i
n
g
t
h
e
l
e
c
t
r
i
c
a
l
a
c
t
i
v
i
t
y
i
n
a
n
o
r
d
e
r
l
y
f
a
s
h
i
o
n
.
T
h
37
e syst
c
o
n
d
u
c
t
i
o
n
s
y
s
t
e
m
c
o
n
s
i
s
t
s
o
f
s
i
n
u
s
n
o
d
e
,
i
n
-
Tab
le
1.6
Con
duc
tion
em
of
the
hea
rt
[2,
6,
9]
Others (525ml)
Heart (215ml)
Skin (490ml)
4%
10%
Abdominal
Kidney (950ml)
19%
10%
organsBrain (650ml)
(1200ml)13%
Skeletal muscle
(1030ml)
20%
24%
39
ter
no
dal
tra
cts
,
A
V
no
de,
A
V
(H
is)
bu
ndl
e,
rig
ht
an
d
lef
t
bu
ndl
e
bra
nc
he
s
an
d
Pu
rki
nje
fib
res
[4,
18
]
(T
abl
e
1.6
).
1
.
3
P
h
y
s
i
o
l
o
g
y
o
f
t
h
e
H
e
a
r
t
1.
3.
1
Ci
rc
ul
at
or
y
Sy
st
e
m:
Sy
st
e
mi
c
a
n
d
P
u
l
m
o
n
a
r
y
C
i
r
c
u
l
a
t
i
o
n
Th
e
car
dio
vas
cul
ar
sys
te
m
del
ive
rs
ox
yg
en
an
d
nut
rie
nts
to
the
tiss
ues
an
d
car
rie
s
aw
ay
wa
ste
ma
teri
als
to
be
eli
mi
nat
ed
by
org
ans
suc
h
as
lun
gs,
liv
er
an
d
kid
ne
ys
[1,
4]
(Fi
g.
1.4
).
Th
is
sys
te
m
is
req
uir
ed
to
fun
cti
on
un
der
var
iou
s
nor
ma
41
lis
ed
an
d
dis
eas
ed
co
ndi
tio
ns.
Th
e
pul
mo
nar
y
an
d
sys
te
mi
c
cir
cul
ati
on
s
tog
eth
er
hel
p
in
ful
fill
ing
thi
s
rol
e.
Pul
mo
nar
y
cir
cul
ati
on
is a
lo
w
res
ist
an
ce,
hig
h
ca
pa
cit
an
ce
be
d,
an
d
sys
te
mi
c
cir
cul
ati
on,
in
co
mp
ari
so
n,
is a
rel
ati
vel
y
hig
h
res
ist
an
ce
vas
cul
ar
be
d
[4,
11,
14,
21]
(Fi
g.
1.5
).
T
he
de
ox
yg
ena
ted
blo
od
fro
m
the
sup
eri
or
ve
na
cav
a
(fr
om
up
per
ext
re
mit
ies,
hea
d,
an
d
che
st
wa
ll),
inf
eri
or
ve
na
cav
a
(tr
un
k,
ab
do
mi
nal
org
ans
an
d
lo
we
r
ext
re
mit
ies
)
an
d
the
cor
on
ary
sin
use
s
(fr
om
my
oca
rdi
um
)
rea
che
s
the
RA
[1,
4,
6,
11]
.
Th
e
RA
is
fill
ed
wit
h
de
ox
yg
ena
ted
blo
od,
inc
rea
sin
g
pre
ssu
re
in
the
atri
al
cha
mb
er.
W
43
he
n
the
atri
al
pre
ssu
re
exc
eed
s
the
pre
ssu
re
in
the
RV
,
the
tric
usp
id
val
ve
op
ens
all
ow
ing
thi
s
blo
od
to
ent
er
the
RV
[1,
4,
6,
11]
.
As
a
res
ult
of
thi
s
fill
ing
,
an
d
as
the
RV
sta
rts
to
co
ntr
act
the
pre
ssu
re
in
the
RV
bui
lds
up
for
cin
g
the
tric
usp
id
val
ve
to
clo
se
an
d
the
pul
mo
nic
val
ve
to
op
en,
the
reb
y
eje
cti
ng
the
blo
od
int
o
the
pul
mo
nar
y
art
eri
es
an
d
lun
gs
[1,
4,
6,
11]
.
T
he
ox
yg
ena
ted
blo
od
fro
m
the
lun
gs
rea
che
s
the
LA
via
the
pul
mo
nar
y
vei
ns
an
d
as
a
res
ult,
pre
ssu
re
in
LA
bui
lds
up
an
d
wh
en
it
exc
eed
s
tha
t of
the
LV,
the
mit
ral
val
ve
op
ens
,
all
ow
ing
the
blo
od
to
ent
er
the
LV
[1,
4,
6,
11]
.
W
he
n
the
blo
od
fill
s
the
LV,
an
d
as
the
LV
sta
rts
to
co
ntr
45
act,
the
LV
cha
mb
er
pre
ssu
re
inc
rea
se
for
ces
the
mit
ral
val
ve
to
clo
se
an
d
aor
tic
val
ve
to
op
en,
thu
s
eje
cti
ng
blo
od
int
o
the
aor
ta,
to
be
dis
trib
ute
d
thr
ou
gh
out
the
bo
dy
[1,
4,
6,
11]
.
1.
3.
2
Co
nd
uc
tio
n
Sy
st
e
m
of
th
e
He
ar
t
(
E
x
c
i
t
a
t
i
o
n
S
e
q
u
e
n
c
e
)
T
h
e
c
a
r
d
i
a
c
m
y
o
c
y
t
e
s
h
a
v
e
g
e
n
e
r
a
t
i
o
n
,
w
h
i
c
h
a
u
n
i
q
u
e
a
b
i
l
i
t
y
o
f
a
u
t
o
m
a
t
i
c
i
m
p
u
l
s
e
r
e
s
u
l
t
s
i
n
a
u
t
o
m
a
t
i
c
r
h
y
t
h
m
i
c
i
t
y
.
N
o
r
47
m
a
l
l
y
t
h
e
t
h
e
e
l
e
c
t
r
i
c
a
l
f
a
s
t
e
s
t
i
m
p
u
l
s
e
b
e
g
i
n
s
i
n
t
h
e
S
A
n
o
d
e
,
h
a
s
i
m
p
u
l
s
e
g
e
n
e
r
a
t
i
o
n
a
b
i
l
i
t
y
a
n
d
a
s
h
e
n
c
e
r
i
v
e
s
t
h
e
r
i
g
h
t
h
e
a
r
t
.
T
h
e
i
m
p
u
l
s
e
t
h
e
n
s
p
r
e
a
d
s
t
o
t
h
e
t
h
e
a
t
r
i
a
l
w
a
l
l
s
d
i
r
e
c
t
l
y
,
t
o
t
h
e
l
e
f
t
r
e
s
t
a
t
r
i
u
m
49
y
t
h
e
i
n
t
e
r
a
t
r
i
a
l
c
o
n
d
u
c
t
i
n
g
f
i
b
r
e
s
a
n
d
t
h
e
n
t
o
t
h
e
o
d
e
(
A
V
j
u
n
c
t
i
o
n
a
l
t
i
s
s
u
e
)
[
6
]
(
F
i
g
.
1
.
6
)
.
T
h
e
A
V
A
V
n
o
d
e
o
n
d
u
c
t
s
t
h
e
i
m
p
u
l
s
e
w
i
t
h
a
d
e
l
a
y
a
n
d
p
r
o
p
a
g
a
t
e
s
t
h
r
o
u
g
h
t
h
e
v
e
n
t
r
i
c
u
l
a
r
m
y
o
c
a
r
d
i
u
m
v
i
a
t
h
e
A
V
b
u
n
d
l
e
o
f
H
i
s
a
n
51
d
P
u
r
k
i
n
j
e
f
i
b
r
e
s
[
1
,
3
,
6
,
8
1
3
,
1
8
]
.
F
r
o
m
t
h
e
P
u
r
k
i
n
j
e
f
i
b
r
e
s
,
t
h
e
e
x
c
i
t
a
t
i
o
n
i
m
p
u
l
s
e
t
h
e
n
c
o
n
t
i
n
u
e
s
t
h
r
o
u
g
h
m
y
o
c
a
r
d
i
a
l
c
e
l
l
s
o
u
t
s
i
d
e
t
h
e
s
p
e
c
i
a
l
i
s
e
d
c
o
n
d
u
c
t
i
o
n
p
a
t
h
w
a
y
t
o
r
e
a
c
h
t
h
e
s
u
b
e
n
d
o
c
a
r
d
i
a
l
s
u
r
f
a
c
e
.
T
h
i
s
r
a
p
i
d
,
s
i
53
m
u
l
t
a
n
e
o
u
s
a
n
d
c
o
o
r
d
i
n
a
t
e
d
s
p
r
e
a
d
o
f
e
x
c
i
t
a
t
i
o
n
t
h
r
o
u
g
h
t
h
e
v
e
n
t
r
i
c
l
e
s
p
r
o
d
u
c
e
s
a
c
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o
r
d
i
n
a
t
e
d
c
o
n
t
r
a
c
t
i
o
n
o
f
b
o
t
h
v
e
n
t
r
i
c
l
e
s
,
t
h
u
s
e
n
s
u
r
i
n
g
e
f
f
i
c
i
e
n
t
p
u
m
p
i
n
g
o
f
b
l
o
o
d
t
o
t
h
e
p
u
l
m
o
n
a
r
y
a
n
d
s
y
s
t
e
m
i
c
c
i
r
c
u
l
a
t
i
o
n
s
[
1
,
3
,
6
,
8
55
1
3
,
a
n
d
1
8
]
.
t
h
e
1
.
3
.
3
A
c
t
i
o
n
P
o
t
e
n
t
i
a
l
(
A
P
)
V
e
n
t
r
i
c
l
e
s
,
a
t
r
i
a
P
u
r
k
i
n
j
e
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y
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t
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m
h
a
v
e
a
s
t
a
b
l
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t
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m
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b
r
a
n
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p
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t
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t
i
a
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o
f
a
b
o
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t
9
0
m
V
,
d
e
t
e
r
m
i
n
e
d
m
a
i
n
l
y
b
y
K
c
t
a
n
c
e
[
1
2
,
1
3
]
.
T
h
e
a
c
t
i
o
n
p
o
t
e
n
t
i
a
l
i
s
o
f
l
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n
d
u
d
u
r
a
t
i
57
o
n
h
e
a
b
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t
a
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t
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o
n
3
0
0
m
s
,
w
h
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c
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i
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y
T
h
e
t
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e
a
c
t
i
o
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a
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p
o
t
e
n
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a
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f
t
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c
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d
i
a
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e
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t
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a
s
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d
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r
a
n
s
i
e
n
t
i
59
n
w
a
r
d
i
n
c
r
e
a
s
e
n
,
r
e
f
e
r
r
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d
t
o
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n
a
s
t
h
e
t
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m
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p
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k
e
c
o
n
d
u
c
t
a
n
c
e
o
f
o
f
t
h
e
N
a
+
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o
t
h
e
a
c
t
i
o
n
p
o
t
e
n
t
i
a
l
o
p
h
a
s
e
b
y
0
(
F
i
g
s
.
1
.
7
a
n
d
1
.
8
)
[
1
2
,
1
3
]
.
T
h
i
s
a
b
r
i
e
f
t
r
a
n
s
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t
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+
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s
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m
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r
61
a
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.
T
h
e
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+
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1
o
f
a
c
t
i
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n
p
o
-
63
u
l
a
t
o
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y
p
a
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w
a
y
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f
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h
e
c
a
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s
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u
l
a
r
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s
t
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m
F
i
g
1
.
5
C
i
r
c
[
8
]
t
e
n
t
i
a
l
[
1
2
,
r
p
h
a
s
e
1
3
]
.
A
f
t
e
r
a
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65
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n
o
u
t
w
a
r
d
K
+
i
o
n
c
o
n
d
u
c
t
a
n
c
e
[
1
2
,
1
3
]
.
T
h
e
s
e
o
u
t
w
a
r
d
a
n
d
i
n
w
a
r
d
c
u
r
r
e
n
t
s
a
r
e
s
u
c
h
t
h
a
t
t
h
e
y
m
a
i
n
t
a
i
n
t
h
e
m
e
m
b
r
a
n
e
p
o
t
e
n
t
i
a
l
i
n
t
h
e
p
l
a
t
e
a
67
u Le
p
h
a
s
e
.
ft
atr
iu
m
an
T
h d
e
the
p
l
a
t
e
a
u
A
V
jun
cti
on
p al
h
a tis
s su
e
e
i
s
f
o
l
l
o
w
e
d
AV no
Bundl
His
b
y
t
h
e
r
e
p
o
-
R
i
g
h
t
a
SA node
n
d
le
ft o
b
uc
n
d
le r
b
r d
a
n
c
h
i
a
e l
s
s
Term
inal
u
Purk
inje
r
fiber
s f
S
u
b
a
c
e
e
F
n
d
i
g
69
1
.
h
e
C
o
t
i
n
g
s
y
s
t
e
m
o
f
lari
sat
ion
ph
ase
of
the
act
ion
pot
ent
ial
or
ph
ase
3,
wh
ich
res
ult
s
fro
m
the
de
cli
nin
g
in
wa
rd
Ca
ion
co
nd
uct
an
ce
an
d
an
inc
rea
se
in
the
out
wa
rd
K+
ion
co
nd
uct
an
ce
[12
,
13]
.
Th
is
out
wa
rd
K+
ion
co
nd
uct
an
ce
hy
per
pol
ari
ses
the
me
mb
ran
e
to
wa
rds
K+
ion
eq
uili
bri
um
an
d
bri
ng
s
ab
out
the
rep
ola
ris
ati
on
or
ph
ase
4
[12
,
13]
.
T
he
SA
no
de
act
ion
pot
ent
ial
is
dif
fer
ent
fro
m
the
res
t of
the
co
nd
uct
ing
sys
te
m
an
d
71
the
my
oca
rdi
um
. It
is
cha
rac
teri
sed
by
an
uns
tab
le
res
tin
g
me
mb
ran
e
pot
ent
ial
or
ph
ase
4.
Thi
s
res
ult
s
fro
m
an
inc
rea
sed
Na
+
ion
co
nd
uct
anc
e
res
ulti
ng
in
in
wa
rd
Na
+
cur
ren
t
[12
,
13]
.
Th
e
in
wa
rd
Na
+
cur
ren
t is
trig
ger
ed
by
the
rep
ola
ris
ati
on
of
the
pre
ced
ing
act
ion
pot
ent
ial.
In
ad
diti
on,
the
ph
ase
0
of
the
act
ion
pot
ent
ial
in
the
SA
no
de
is
the
res
ult
of
in
wa
rd
Ca+
+
ion
co
nd
uct
anc
e
in
co
ntr
ast
to
Na
+
ion
, as
in
the
res
t of
the
my
oca
rdi
um
[12
,
13]
. In
ad
diti
on,
the
SA
no
de
act
ion
pot
ent
ial
lac
ks
the
pla
tea
u
or
ph
ase
2
of
the
my
oca
rdi
al
cel
l
act
ion
pot
ent
ial
[12
,
13]
.
Th
e
ups
tro
ke
of
the
act
ion
pot
ent
ial
in
the
AV
no
de
is
als
o
du
e
to
the
Ca+
+
ion
co
nd
uct
+30Phase 1.
+20K efflux
+100
0- Gnanasegaran,
Phase and
2 John R Buscombe
Syed Shah, Gopinath
Jeanette Sundberg-Cohon,
-10Ca influx, K efflux.
-20Phase 0
15Phase 0
Na influx
Ca influx Phase 3
-50Phase
K efflux 3
30-60Phase 4
K efflux.
-70Resting membrane potential.
-8045-90Fig 1.7 Cardiac action potential
15-
73
6075-
Phase 4
Na influx
anc
e
as
in
the
SA
no
de.
Th
e
co
nd
uct
ion
vel
oci
ty
is
fas
test
in
the
Pur
kin
je
sys
te
m
an
d
slo
we
st
in
the
AV
no
de
[12
,
13]
.
1.
3.
4
M
ec
ha
ni
s
m
of
Ex
cit
ati
on
an
d
Co
nt
ra
cti
on
C
o
u
p
l
i
n
g
o
f
C
a
r
d
i
a
c
M
y
o
c
y
t
e
s
Th
e
reg
ula
tio
n
of
car
dia
c
mu
scl
e
co
ntr
act
ion
has
ne
ura
l,
hor
mo
nal
an
d
int
rin
sic
co
mp
on
ent
s.
Th
e
he
art
is
co
mp
ose
d
of
cyl
ind
ric
al
car
dia
c
cel
ls
an
d
sti
mu
lati
on
of
on
e
car
dia
c
cel
l
init
iat
es
sti
mu
lati
on
of
adj
ace
nt
cel
ls
[7,
75
22]
. In
ge
ner
al,
the
car
dia
c
cel
ls
are
of
tw
o
typ
es
(a)
ele
ctri
cal
cel
ls
an
d
(b)
my
oc
ard
ial
cel
ls
[7]
.
Th
e
ele
ctri
cal
cel
ls
are
spe
cia
lis
ed
my
oc
ard
ial
cel
ls
tha
t
ha
ve
ess
ent
iall
y
los
t
the
abi
lity
to
co
ntr
act
but
ha
ve
be
co
me
spe
cia
lis
ed
in
the
co
nd
uct
ion
of
car
dia
c
im
pul
ses
.
T
he
my
oca
rdi
al
cel
ls
ha
ve
tw
o
spe
cifi
c
pro
per
ties
:
co
ntr
act
ilit
y
(ab
ilit
y
of
the
cel
ls
to
sho
rte
n
an
d
ret
urn
to
the
ir
ori
gin
al
len
gth
)
an
d
ext
ens
ibil
ity
(ce
ll
fila
me
nts
abi
lity
to
str
etc
h)
[22
].
Th
e
co
ntr
act
ion
me
cha
nis
m
co
nta
ins
nu
me
rou
s
ste
ps
an
d
ex
cit
ati
on
co
ntr
act
ion
co
upl
ing
is
the
ter
m
use
d
to
def
ine
the
eve
nts
tha
t
tra
nsl
ate
the
de
pol
ari
sati
on
of
the
car
dia
c
77
cel
l
me
mb
ran
e
to
the
co
ntr
act
ion
of
the
mu
scl
e
fib
res
[5,
6,
11]
(Fi
g.
1.9
).
Th
e
co
ntr
act
ile
fila
me
nts
act
in
an
d
my
osi
n
in
the
my
oca
rdi
um
are
res
po
nsi
ble
for
the
co
ntr
act
ion
. In
the
car
dia
c
cel
l,
the
sar
co
pla
sm
ic
reti
cul
um
an
d
the
cist
ern
ae
co
nta
ins
hig
h
co
nce
ntr
ati
ons
of
ion
ise
d
Ca+
+
an
d
de
pol
ari
sati
on
of
the
T
tub
ule
s,
wh
ich
is
an
ext
ens
ion
of
the
cel
l
me
mb
ran
e,
cau
ses
the
rel
eas
e
of
Ca+
+
fro
m
the
se
str
uct
ure
s
[5,
6,
11]
. In
ge
ner
al,
an
act
ion
pot
ent
ial
pre
ced
es
eac
h
co
ntr
act
ion
,
an
d
bet
we
en
co
ntr
act
ion
s
the
Ca+
+
wit
hin
the
cel
l is
ver
y
lo
w.
Th
e
car
dia
c
act
ion
pot
ent
ial
is
uni
qu
e
in
tha
t it
has
a
pla
tea
u
ph
ase
,
wh
ich
is
ma
int
ain
ed
by
the
in
79
wa
rd
Ca+
+
infl
ux
(cu
rre
nt)
[5,
6,
11]
. It
is
thi
s
in
wa
rd
Ca+
+
cur
ren
t
tha
t
trig
ger
s
rel
eas
e
of
fur
the
r
Ca+
+
fro
m
the
sar
co
pla
sm
ic
reti
cul
um
.
Th
e
am
ou
nt
of
Ca+
+
rel
eas
ed
de
pe
nds
on
the
str
en
gth
of
the
in
wa
rd
Ca+
+
cur
ren
t.
Co
mp
are
d
wit
h
the
res
tin
g
or
bas
al
co
ndi
tio
ns,
the
co
nce
ntr
ati
on
of
Ca+
+
is
ten
tim
es
hig
her
dur
ing
an
AP
[5]
.
W
he
n
the
co
nce
ntr
ati
on
of
the
Ca+
+
is
hig
h,
the
tro
po
nin
-C
is
bo
un
d
by
fou
r
Ca+
+
ion
s
per
mo
lec
ule
an
d
thi
s
cha
ng
es
the
sha
pe
of
tro
po
nin
[6,
7,
11]
.
Thi
s
cha
ng
e
in
sha
pe
all
ow
s
the
tro
po
my
osi
n
to
un
co
ver
the
cro
ssbri
dg
e
site
s,
wh
ich
all
ow
s
the
for
ma
tio
n
of
cro
ss
bri
dg
es
bet
we
en
act
in
an
d
my
81
osi
n
fila
me
nts
[6,
7,
11]
. In
the
res
tin
g
stat
e,
the
tro
po
nin
mo
lec
ule
s
shi
eld
the
cro
ssbri
dg
es.
T
he
pre
sen
ce
of
ad
en
osi
ne
tri
ph
os
ph
ate
(A
TP
) is
im
por
tan
t in
the
co
ntr
act
ile
pro
ces
s.
Th
e
a
c
t
i
n
m
y
o
s
i
n
c
r
o
s
s
b
r
i
d
g
e
s
a
r
e
f
o
r
m
e
d
i
n
t
h
e
p
r
e
s
e
n
c
e
o
f
A
T
P
,
w
h
i
c
h
l
a
t
e
r
h
y
d
r
o
l
y
s
e
s
.
T
h
e
e
n
e
r
g
y
d
e
r
i
v
e
d
f
r
o
83
m
A
T
P
h
y
d
r
o
l
y
s
i
s
l
e
a
d
s
t
o
c
h
a
n
g
e
s
i
n
t
h
e
m
y
o
s
i
n
f
i
g
u
r
a
t
i
o
n
s
o
t
h
a
t
a
c
t
i
n
c
a
n
b
e
d
e
t
a
c
h
e
d
f
r
o
m
h
e
a
d
m
y
o
s
i
n
c
o
n
(
1
0
n
m
/
c
y
c
l
e
)
.
F
i
n
a
l
l
y
,
A
T
P
i
s
r
e
q
u
i
r
e
d
f
o
r
r
e
l
a
x
a
t
i
o
n
o
f
t
h
e
m
u
s
c
l
e
b
y
d
i
s
s
o
c
i
a
t
i
n
g
t
h
e
c
r
o
s
s
b
r
i
d
g
e
s
[
6
]
.
I
n
g
e
85
n
e
r
a
l
,
c
r
e
a
s
e
a
n
i
n
i
n
c
r
e
a
s
e
t
h
e
i
n
t
h
e
n
u
m
b
e
r
o
f
b
r
i
d
g
e
s
c
a
u
s
e
s
a
n
i
n
f
o
r
c
e
o
f
c
o
n
t
r
a
c
t
i
o
n
,
w
h
i
c
h
i
s
p
r
o
p
o
r
t
i
o
n
a
l
t
o
t
h
e
i
n
t
r
a
c
e
l
l
u
l
a
r
C
a
+
+
c
o
n
c
e
n
t
r
a
t
i
o
n
[
6
]
.
R
e
l
a
x
a
t
i
o
n
o
c
c
u
r
s
w
h
e
n
t
h
e
i
n
t
r
a
c
e
l
l
u
l
a
r
C
a
+
+
i
s
r
e
s
e
q
u
e
s
t
e
r
e
d
87
b
a
c
k
i
n
t
o
t
h
e
s
a
r
c
o
p
l
a
s
m
i
c
r
e
t
i
c
u
l
u
m
b
y
t
h
e
C
a
+
+
A
T
P
a
s
e
p
u
m
p
.
A
d
m
i
n
i
s
t
r
a
t
i
o
n
o
f
d
r
u
g
s
l
i
k
e
e
p
i
n
e
p
h
r
i
n
e
,
n
o
r
e
p
i
n
e
p
h
r
i
n
e
,
a
n
d
d
i
g
i
t
a
l
i
s
a
n
d
s
y
m
p
a
t
h
e
t
i
c
n
e
r
v
e
s
t
i
m
u
l
a
t
i
o
n
i
n
c
r
e
a
s
e
s
t
h
e
i
n
t
r
a
c
e
l
l
u
l
a
r
C
a
+
+
l
e
v
e
l
s
,
w
h
i
c
h
r
e
s
u
l
t
89
s
i
n
f
o
r
c
e
f
u
l
c
o
n
t
r
a
c
t
i
o
n
[
6
]
.
N
o
r
e
p
i
n
e
p
h
r
i
n
e
n
o
t
o
n
l
y
c
a
u
s
e
s
f
o
r
c
e
f
u
l
c
o
n
t
r
a
c
t
i
o
n
b
u
t
a
l
s
o
s
h
o
r
t
e
n
s
t
h
e
A
P
a
n
d
c
o
n
t
r
a
c
t
i
o
n
b
y
c
a
u
s
i
n
g
m
o
r
e
r
a
p
i
d
u
p
t
a
k
e
o
f
C
a
s
a
r
c
o
p
l
a
s
m
i
c
r
e
t
i
c
u
l
u
m
[
6
]
.
T
h
e
c
o
n
t
r
a
c
t
i
l
i
t
y
+
+
b
y
t
h
e
c
a
n
b
e
q
91
u
a
n
t
i
f
i
e
d
b
y
c
a
l
c
u
l
a
t
i
n
g
t
h
e
e
j
e
c
t
i
o
n
f
r
a
c
t
i
o
n
(
E
F
)
,
w
h
i
c
h
i
s
d
e
f
i
n
e
d
a
s
t
h
e
r
a
t
i
o
o
f
s
t
r
o
k
e
v
o
l
u
m
e
(
S
V
)
t
o
e
n
d
-
d
i
a
s
t
o
l
i
c
v
o
l
u
m
e
(
E
D
V
)
e
l
e
c
t
r
i
c
a
l
c
e
l
l
s
a
r
e
s
p
e
c
i
a
l
S
V
/
E
D
V
)
c
a
r
d
i
a
c
[
1
1
,
c
e
l
l
s
(
E
F
2
3
,
2
4
]
.
T
h
e
o
f
t
h
e
c
o
n
d
u
93
c
t
i
n
g
s
y
s
t
e
m
[
2
2
]
.
T
h
e
s
e
c
e
l
l
s
a
r
e
p
r
i
m
a
r
i
l
y
r
e
s
p
o
n
s
i
b
l
e
f
o
r
t
h
e
f
o
r
m
a
t
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[
7
]
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[
7
,
2
2
]
.
I
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p
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,
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107
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109
+
-
Na+
K+
Resting state
Na+ Beginning of
depolarisation
+
-
+
-
K+
Completion of depolarisation
+
+
Na
Na+
Beginning of repolarisation
K+
Completion of repolarisation
+
K+
Na+
reg
ula
ted
by
ele
ctr
ica
l,
me
ch
ani
cal
or
ch
em
ica
l
sti
mu
li
[7]
.
Fu
rth
er
mo
re,
the
co
nc
ent
rat
ion
gra
die
nt
aff
ect
s
the
ion
dis
tri
but
ion
acr
oss
the
cel
l
me
mb
ran
e
[7]
.
I
n
the
res
tin
g
stat
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of
the
cel
l,
the
ele
ctri
cal
act
ivit
y
is
mo
re
pos
itiv
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out
sid
e
the
cel
l
an
d
mo
re
ne
gat
ive
ins
ide
the
cel
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an
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no
ele
ctri
cal
act
ivit
y
occ
urs
[7,
22]
.
W
he
n
the
cel
111
l is
sti
mu
lat
ed,
the
per
me
abi
lity
of
the
me
mb
ran
e
cha
ng
es
all
ow
ing
sod
iu
m
to
ent
er
the
cel
l
[7]
.
Thi
s
res
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s
in
the
ins
ide
of
the
cel
l
bec
om
ing
mo
re
pos
itiv
e
tha
n
the
out
sid
e,
res
ulti
ng
in
a
de
pol
ari
sed
stat
e
[7]
.
On
ce
de
pol
ari
sati
on
is
co
mp
let
e,
the
me
mb
ran
e
all
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sod
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m
to
exi
t,
on
ce
aga
in
ma
kin
g
the
cel
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mo
re
ne
1
.
Fig 1.11Phases of cardiac cycle
gat
Atrial
Systole
Passive
ventricular
filling
Isovolumetric
ventricular
Contraction
Phases of
Cardiac cycle
Isovolumetric
ventricular
relaxation
Ejection
ive
ins
ide
.
Thi
s
en
d
res
ult
is
cal
led
rep
ola
ris
ati
on
(ce
ll
rec
ov
ery
)
[7]
.
3
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5
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m
u
s
c
l
e
b
u
t
a
n
d
t
h
e
y
a
r
e
i
n
v
o
l
v
e
d
i
n
r
e
g
u
l
a
t
i
o
n
o
f
b
l
o
o
d
f
l
o
w
a
n
d
s
y
s
t
e
m
i
c
b
l
o
o
d
p
r
e
s
s
u
r
e
[
4
6
,
8
,
1
1
1
4
]
.
T
h
e
c
h
o
l
i
n
e
r
g
i
c
o
r
t
h
e
p
a
r
a
s
y
m
p
125
a
t
h
e
t
i
c
n
e
r
v
o
u
s
s
y
s
t
e
m
e
f
f
e
c
t
s
i
n
t
h
e
h
e
a
r
t
a
r
e
o
p
p
o
s
i
t
e
t
o
t
h
e
o
n
e
s
m
e
n
t
i
o
n
e
d
a
b
o
v
e
a
n
d
t
h
e
t
r
a
n
s
m
i
t
t
e
r
i
n
v
o
l
v
e
d
i
s
m
a
i
n
l
y
a
c
e
t
y
l
c
h
o
l
i
n
e
[
4
6
,
8
,
1
1
1
4
]
.
T
h
e
p
l
y
t
h
e
p
a
r
a
s
y
m
p
a
t
h
e
t
i
c
f
i
b
r
e
s
t
o
t
h
e
h
e
a
r
t
v
i
a
v
a
g
i
t
h
e
s
u
p
c
a
r
127
d
i
a
c
p
l
e
x
u
s
e
s
.
T
h
e
p
a
r
a
s
y
m
p
a
t
h
e
t
i
c
e
f
f
e
c
t
s
a
r
e
m
e
d
i
a
t
e
d
v
i
a
t
h
e
m
u
s
c
a
r
a
n
i
c
r
e
c
e
p
t
o
r
s
,
w
h
i
c
h
a
c
t
b
y
i
n
h
i
b
i
t
i
o
n
o
f
a
d
e
n
y
l
a
t
e
c
y
c
l
a
s
e
a
n
d
h
e
n
c
e
c
A
M
P
l
e
v
e
l
s
a
n
d
r
e
s
u
l
t
i
n
a
d
e
c
r
e
a
s
e
d
e
c
r
e
a
s
e
t
h
e
i
n
i
n
t
r
a
c
e
l
l
u
l
a
r
h
e
a
r
t
r
a
t
e
,
c
o
129
n
t
r
a
c
t
i
o
n
a
n
d
c
o
n
d
u
c
t
i
o
n
v
e
l
o
c
i
t
y
[
4
6
,
8
,
1
1
1
4
]
.
T
he
aut
on
om
ic
cen
tre
s
in
the
CN
S,
ma
inl
y
the
vas
om
oto
r
cen
tre
of
me
dul
la
an
d
the
hy
pot
hal
am
us
reg
ula
te
the
bal
anc
e
bet
we
en
the
lev
el
of
sy
mp
ath
eti
c
an
d
the
par
asy
mp
ath
eti
c
out
put
to
the
car
dio
vas
cul
ar
sys
te
m
de
pe
ndi
ng
on
the
aff
ere
nt
inp
uts
fro
m
the
per
iph
ery
an
d
the
CN
S
[4
6,
8,
11
14]
.
Th
ere
is
nor
ma
lly
a
ton
ic
va
gal
dis
cha
rge
in
hu
ma
ns
wh
ich
ov
erri
des
the
mo
der
ate
ton
ic
dis
cha
rge
in
the
car
dia
c
sy
mp
ath
eti
c
ner
ves
[12
,13
].
Bo
th
the
sy
mp
ath
eti
c
an
d
the
par
asy
mp
ath
eti
c
fib
res
in
131
the
spl
anc
hni
c
tho
rac
ic
ner
ves
an
d
the
va
gi
car
ry
aff
ere
nt
inp
ut
me
dia
ted
via
bar
ore
cep
tor
s
an
d
che
mo
rec
ept
ors
to
the
aut
on
om
ic
cen
tre
s
in
the
CN
S,
in
ad
diti
on
to
the
eff
ere
nt
out
put
fro
m
the
CN
S
[12
,
13]
.
Th
ese
aff
ere
nts
an
d
eff
ere
nts
are
inv
olv
ed
in
me
dia
tio
n
of
car
dio
vas
cul
ar
refl
exe
s
suc
h
as
bar
ore
cep
tor
an
d
the
che
mo
rec
ept
or
refl
exe
s
[4
6,
8,
11
14]
.
1
.
3
.
6
C
a
r
d
i
a
c
C
y
c
l
e
Th
e
car
dia
c
cy
cle
co
nsi
sts
of
co
ntr
act
ion
an
d
rel
ax
ati
on
of
bot
h
atri
a
an
d
ve
ntr
icl
es
(Fi
g.
1.1
1).
Th
e
he
art
do
es
not
co
ntr
act
or
rel
ax
as
a
sin
gle
uni
t
[11
].
Th
e
car
dia
c
cy
cle
co
nsi
sts
of
cy
cli
c
str
uct
ura
l
133
an
d
fun
cti
on
al
ch
an
ges
wh
ich
the
he
art
un
der
go
es
ev
ery
0.8
s
on
av
era
ge
to
ma
int
ain
the
blo
od
flo
w
thr
ou
gh
the
bo
dy.
Th
e
cy
cle
sta
rts
wit
h
an
atri
al
sys
tol
e,
wh
ich
is
pre
ce
de
d
by
an
im
pul
se
ge
ner
ate
d
in
the
SA
no
de
an
d
is
spr
ea
d
acr
oss
the
atri
a
cor
res
po
ndi
ng
to
a
Pwa
ve
on
the
ele
ctr
oc
ard
iog
ra
m
[11
].
Th
e
atri
al
sys
tol
e
co
ntr
ibu
tes
to,
but
is
not
ess
ent
ial
for
,
ve
ntr
icu
lar
fill
ing
,
but
it
do
es
be
co
me
an
im
por
tan
t
fac
tor
in
a
dis
eas
ed
he
art
[11
,
14]
.
Fil
lin
g
of
the
ve
ntr
icl
es
by
atri
al
co
ntr
act
ion
ca
use
s
the
fou
rth
he
art
so
un
d,
thi
s is
not
nor
ma
lly
au
dib
le
in
ad
ult
s.
Du
rin
g
thi
s
ph
ase
,
the
AV
val
ves
are
op
en
an
d
the
ve
ntr
icl
es
135
are
rel
axi
ng.
Th
e
se
mil
un
ar
val
ves
(S
L)
are
clo
sed
,
wh
ich
pre
ve
nts
reent
ry
of
blo
od
fro
m
the
pul
mo
nar
y
art
ery
or
the
aor
ta.
Th
e
iso
vol
um
etri
c
ve
ntr
icu
lar
co
ntr
act
ion
(st
art
of
ve
ntr
icu
lar
sys
tol
e
wh
ile
the
SL
val
ves
are
clo
sed
)
fol
lo
ws
the
atri
al
sys
tol
e,
dur
ing
wh
ich
the
ve
ntr
icu
lar
vol
um
e
re
ma
ins
co
nst
ant
[6,
11,
14]
.
Th
e
iso
vol
um
etri
c
ve
ntr
icu
lar
co
ntr
act
ion
is
pre
ce
de
d
by
the
spr
ea
d
of
the
ele
ctri
c
im
pul
se
fro
m
the
AV
no
de
to
the
res
t of
the
ve
ntr
icu
lar
my
oc
ard
iu
m,
an
d
thi
s is
see
n
as
the
Q
RS
co
mp
lex
on
an
ele
ctr
oc
ard
iog
ra
m
[6,
11,
14]
.
W
he
n
the
ve
ntr
icu
lar
pre
ssu
re
be
co
me
s
gre
ate
r
tha
n
the
atri
al
pre
ssu
re,
the
AV
val
ves
clo
se,
cor
res
po
137
ndi
ng
to
the
firs
t
he
art
so
un
d,
wit
h
the
mit
ral
val
ve
clo
sin
g
fra
cti
on
all
y
bef
ore
the
tric
us
pid
val
ve.
Th
e
se
mil
un
ar
(S
L)
val
ves
op
en
as
the
pre
ssu
re
in
the
ve
ntr
icl
es
ex
cee
ds
tha
t of
the
pre
ssu
re
in
the
aor
ta
an
d
the
pul
mo
nar
y
art
ery
,
res
ulti
ng
in
rap
id
ve
ntr
icu
lar
eje
cti
on.
Th
is
res
ult
s
in
a
dra
ma
tic
red
uct
ion
in
the
ve
ntr
icu
lar
vol
um
e
as
mo
st
of
the
ve
ntr
icu
lar
blo
od
(str
ok
e
vol
um
e)
is
eje
cte
d
dur
ing
thi
s
ph
ase
.
Th
e
atri
a
als
o
be
gin
to
fill
dur
ing
thi
s
ph
ase
.
Th
e
en
d
of
ve
ntr
icu
lar
co
ntr
act
ion
cor
res
po
nd
s
to
the
on
set
of
the
Twa
ve
on
the
ele
ctr
oc
ard
iog
ra
m
[6,
11,
14]
.
To
wa
rds
the
en
d
of
the
ve
ntr
icu
lar
co
ntr
act
ion
the
eje
cti
on
of
blo
139
od
fro
m
the
ve
ntr
icl
es
is
slo
we
r
as
the
ve
ntr
icu
lar
pre
ssu
re
sta
rts
to
dro
p.
Th
is
is
fol
lo
we
d
by
iso
vol
um
etri
c
ve
ntr
icu
lar
rel
ax
ati
on
ma
rke
d
by
clo
sur
e
of
the
aor
tic
an
d
pul
mo
nic
val
ves
.
Th
e
clo
sur
e
of
the
se
mil
un
ar
val
ves
cor
res
po
nd
s
to
the
sec
on
d
he
art
so
un
d.
Th
e
AV
val
ves
are
clo
sed
an
d
the
ve
ntr
icu
lar
vol
um
e
re
ma
ins
co
nst
ant
.
Th
e
pre
ssu
re
in
the
ve
ntr
icl
es
is
red
uc
ed
rap
idl
y
[6,
11,
14]
.
W
he
n
the
ve
ntr
icu
lar
pre
ssu
re
be
co
me
s
les
s
tha
n
the
atri
al
pre
ssu
re,
the
AV
val
ves
op
en
ma
rki
ng
the
en
d
of
the
iso
vol
um
etri
c
ph
ase
of
ve
ntr
icu
lar
rel
ax
ati
on
an
d
the
on
set
of
rap
id
ve
ntr
icu
lar
fill
ing
.
Du
rin
g
the
be
gin
nin
g
of
thi
s
141
ph
ase
of
ve
ntr
icu
lar
rel
ax
ati
on,
the
re
is
pas
siv
e
fill
ing
of
the
ve
ntr
icl
es
wit
h
the
blo
od
fro
m
the
atri
a,
wh
ich
is
rap
id
[6,
11,
14]
.
Th
e
rap
id
flo
w
of
blo
od
fro
m
the
atri
a
int
o
ve
ntr
icl
es
ca
use
s
the
thi
rd
he
art
so
un
d,
wh
ich
is
nor
ma
l in
chi
ldr
en
but
in
ad
ult
s is
ass
oci
ate
d
wit
h
dis
eas
e.
Re
du
ce
d
ve
ntr
icu
lar
fill
ing
so
me
tim
es
ref
err
ed
to
as
dia
sta
sis,
wh
ich
is
the
lon
ges
t
per
iod
of
the
car
dia
c
cy
cle
,
fol
lo
ws
the
rap
id
fill
ing
.
M
ost
of
the
my
oc
ard
iu
m
rec
eiv
es
its
blo
od
su
ppl
y
dur
ing
thi
s
ph
ase
of
the
car
dia
c
cy
cle
[6,
11,
14]
.
143
1
.
3
.
7
P
h
y
s
i
o
l
o
g
y
o
f
C
o
r
o
n
a
r
y
C
i
r
c
u
l
a
t
i
o
n
T
h
e
p
r
i
m
a
r
y
f
u
n
c
t
i
o
n
o
f
t
h
e
c
o
r
o
n
a
r
y
c
i
r
c
u
l
a
t
i
o
n
i
s
t
o
m
e
e
t
t
h
e
m
e
t
a
b
o
l
i
c
d
e
m
a
n
d
o
f
t
h
e
h
e
a
r
t
.
C
o
r
o
n
a
r
y
b
l
o
o
d
f
l
o
w
i
n
c
r
e
a
s
e
s
f
r
o
m
t
h
e
b
a
s
e
l
i
n
e
o
r
r
e
s
t
i
n
g
l
e
v
e
l
t
o
t
h
e
m
a
x
i
m
u
m
d
e
p
e
n
d
145
i
n
g
o
n
m
y
o
c
a
r
d
i
a
l
o
x
y
g
e
n
r
e
q
u
i
r
e
m
e
n
t
s
t
e
i
n
c
r
e
a
s
e
o
f
c
o
r
o
n
a
r
y
b
l
o
o
d
f
l
o
w
i
s
[
2
5
2
7
]
.
r
e
q
u
i
r
e
d
A
n
t
o
a
d
e
q
u
a
m
e
e
t
m
y
o
c
a
r
d
i
a
l
o
x
y
g
e
n
c
o
n
s
u
m
p
t
i
o
n
.
D
u
r
i
n
g
s
t
r
e
n
u
o
u
s
e
x
e
r
c
i
s
e
o
r
m
e
t
a
b
o
l
i
c
d
e
m
a
n
d
,
t
h
e
c
o
r
o
n
a
r
y
b
l
o
o
d
f
l
o
w
i
n
c
r
e
a
s
e
s
147
u
p
t
o
4
6
t
i
m
e
s
[
2
5
2
7
]
.
I
n
c
o
n
d
i
t
i
o
n
s
s
u
c
h
a
s
l
e
f
t
v
e
n
t
r
i
c
u
l
a
r
h
y
p
e
r
t
r
o
p
h
y
(
L
V
H
)
,
m
y
o
c
a
r
d
i
a
l
i
s
c
h
a
e
m
i
a
,
a
n
d
d
i
a
b
e
t
e
s
o
w
m
e
l
l
i
t
u
s
b
e
(
D
M
)
t
h
e
n
o
r
m
a
l
i
n
c
r
e
a
s
e
i
n
c
a
n
b
l
u
n
t
e
d
.
T
h
e
e
p
i
c
a
r
d
i
a
l
c
o
n
d
u
i
t
c
o
r
o
n
a
r
y
r
e
s
i
s
t
a
n
c
e
f
l
i
s
149
p
r
o
d
u
c
e
d
b
y
a
t
h
e
r
o
s
c
l
e
r
o
t
i
c
s
t
e
n
o
s
i
s
a
n
d
t
h
e
d
i
s
t
a
l
v
a
s
c
u
l
a
r
b
e
d
m
a
i
n
t
a
i
n
s
s
a
t
i
s
f
a
c
t
o
r
y
b
l
o
o
d
f
l
o
w
b
y
d
i
l
a
t
i
n
g
.
T
h
e
m
a
x
i
m
a
l
i
n
c
r
e
a
s
e
i
n
c
o
r
o
n
a
r
y
f
l
o
w
a
b
o
v
e
r
e
s
t
i
n
g
l
e
v
e
l
s
i
s
d
e
f
i
n
e
d
a
s
c
o
r
o
n
a
r
y
f
l
o
w
r
e
s
e
r
v
e
(
C
R
F
)
[
2
7
,
2
8
]
.
151
T
h
e
a
r
e
m
a
j
o
r
h
e
a
r
t
d
e
t
e
r
m
i
n
a
n
t
s
r
a
t
e
o
f
m
y
o
c
a
r
d
i
a
l
o
x
y
g
e
n
c
o
n
s
u
m
p
t
i
o
n
(
H
R
)
,
m
y
o
c
a
r
d
i
a
l
w
a
l
l
t
e
n
s
i
o
n
/
s
t
r
e
s
s
(
a
f
t
e
r
l
o
a
d
)
a
n
d
i
n
o
t
r
o
p
i
c
s
t
a
t
e
(
c
o
n
t
r
a
c
t
i
l
i
t
y
)
o
f
o
c
a
r
d
i
u
m
[
2
7
,
2
9
]
.
V
a
s
c
u
l
a
r
r
e
s
i
s
t
a
n
c
e
a
l
s
o
p
l
a
y
s
t
h
e
a
n
m
y
i
m
153
p
o
r
t
a
n
t
r
o
l
e
i
n
c
o
r
o
n
a
r
y
c
i
r
c
u
l
a
t
i
o
n
a
n
d
i
s
m
a
i
n
l
y
d
e
t
e
r
m
i
n
e
d
b
y
m
y
o
c
a
r
d
i
a
l
o
x
y
g
e
n
c
o
n
s
u
m
p
t
i
o
n
,
a
n
d
m
o
d
u
l
a
t
e
d
m
a
i
n
l
y
w
i
t
h
b
y
s
o
m
e
l
o
c
a
l
m
e
t
a
b
o
l
i
c
f
a
c
t
o
r
s
(
v
a
s
o
a
c
t
i
v
e
a
u
t
a
c
o
i
d
s
)
c
o
n
t
r
i
b
u
t
i
o
n
f
r
o
m
n
e
u
r
a
l
s
t
i
m
u
l
i
,
a
n
d
c
i
r
c
u
l
a
155
t
i
n
g
v
a
s
o
a
c
t
i
v
e
s
u
b
s
t
a
n
c
e
s
[
2
7
]
.
1
.
3
.
8
C
o
r
o
n
a
r
y
C
o
l
l
a
t
e
r
a
l
s
T
h
e
i
n
i
t
i
a
l
d
o
c
u
m
e
n
t
a
t
i
o
n
o
f
c
o
r
o
n
a
r
y
c
o
l
l
a
t
e
r
a
l
s
i
n
h
u
m
a
n
s
w
a
s
n
o
t
m
a
d
e
u
n
t
i
l
1
9
6
4
,
a
l
t
h
o
u
g
h
t
h
e
i
r
p
r
e
s
e
n
c
e
w
a
s
c
o
n
s
i
d
e
r
e
d
a
s
f
a
r
b
a
c
k
a
s
157
t
h
e
1
6
t
h
c
e
n
t
u
r
y
[
2
1
,
3
0
]
.
T
h
e
r
e
a
r
e
f
e
w
t
e
r
a
l
v
e
s
s
e
l
s
a
n
d
t
h
e
y
m
a
y
d
e
v
e
l
o
p
i
n
t
o
a
a
n
d
s
m
a
l
l
c
o
l
l
a
m
a
j
o
r
v
a
s
c
u
l
a
r
n
e
t
w
o
r
k
i
n
p
a
t
i
e
n
t
s
w
i
t
h
o
b
s
t
r
u
c
t
i
v
e
i
s
e
a
s
e
[
2
1
]
.
I
t
i
s
g
e
n
e
r
a
l
l
y
b
e
l
i
e
v
e
d
c
o
r
o
n
a
r
y
t
h
a
t
a
r
t
e
r
y
c
o
l
l
a
t
e
r
t
h
e
159
a
l
v
e
s
s
e
l
s
,
o
r
n
e
t
w
o
r
k
,
d
e
v
e
l
o
p
o
n
l
y
w
h
e
n
t
h
e
o
c
c
l
u
s
i
v
e
d
i
s
e
a
s
e
o
r
s
t
e
n
o
s
i
s
i
s
s
e
v
e
r
e
e
n
o
u
g
h
t
o
p
r
o
d
u
c
e
a
s
u
b
s
t
a
n
t
i
a
l
t
r
a
n
s
s
t
e
n
o
t
i
c
p
r
e
s
e
n
t
p
r
e
s
s
u
r
e
d
r
o
p
[
2
1
]
.
W
h
e
n
c
o
l
l
a
t
e
r
a
l
a
r
e
i
n
p
a
t
i
e
n
t
s
w
i
t
h
s
e
v
e
r
e
o
r
s
i
g
n
i
f
i
c
a
n
t
o
c
161
c
l
u
s
i
v
e
C
A
D
,
t
h
e
s
t
e
n
o
t
i
c
v
e
s
s
e
l
p
a
r
t
s
e
e
m
s
o
f
t
o
t
h
e
h
a
v
e
m
y
o
c
a
r
d
i
u
m
s
u
p
p
l
i
e
d
b
y
t
h
e
b
e
t
t
e
r
c
o
n
t
r
a
c
t
i
l
e
f
u
n
c
t
i
o
t
h
a
n
i
s
i
n
p
a
t
i
e
n
t
s
w
i
t
h
o
u
t
c
o
l
l
a
t
e
r
a
l
s
[
3
1
]
.
F
u
r
t
h
e
r
,
i
h
y
p
o
t
h
e
s
i
s
e
d
t
h
a
t
p
r
e
e
x
i
s
t
i
n
g
c
o
l
l
a
t
e
r
a
l
s
m
a
y
a
l
s
163
o
p
l
a
y
a
n
i
m
p
o
r
t
a
n
t
r
o
l
e
b
y
d
e
c
r
e
a
s
i
n
g
t
h
e
e
x
t
e
n
t
a
n
d
d
e
g
r
e
e
o
f
m
y
o
c
a
r
d
i
a
l
d
a
m
a
g
e
a
t
t
h
e
t
i
m
e
o
r
d
u
r
i
n
g
a
n
a
c
u
t
e
c
o
r
o
n
a
r
y
e
p
i
s
o
d
e
[
2
1
]
.
I
n
g
e
n
e
r
a
l
,
t
h
e
cor
on
ary
col
lat
era
ls
are
ca
pa
ble
of
pre
ser
vin
g
the
fu
nct
ion
an
d
str
uct
ure
in
Table 1.7 Physiological changes and cardiovascular response to moderate exercise [33, 3538]
Fatigue
Muscle fatigue
Acidosis
Hyperthermia
CO increases
Dehydration
Increased NH3
SV increases
Hypoglycaemia
Electrochemical changes
EDV increases
Muscle damage
Electrolyte imbalance
ATP Reduction
Increased ADP
TPR decreases
Oxygen extraction increases
Blood flow to heart, muscle and
skin increases
Blood flow to brain increases (slightly)
Blood flow to other viscera decreases
165
a
res
tin
g
my
oc
ard
iu
m,
ho
we
ver
,
the
cor
on
ary
flo
w
per
gra
m
in
a
col
lat
era
lde
pe
nd
ent
my
oc
ard
iu
m
is
red
uc
ed
wh
en
co
mp
are
d
wit
h
the
no
rm
all
y
per
fus
e
d
m
y
o
c
a
r
d
i
u
m
.
O
n
e
r
e
a
s
o
n
f
o
r
t
h
i
s
c
o
u
l
d
b
e
t
h
e
c
h
a
n
g
e
i
n
p
r
e
s
s
u
r
e
g
r
a
d
i
e
n
t
a
c
r
o
s
s
t
h
e
c
o
l
l
a
t
e
r
a
l
v
e
s
s
e
l
s
(
a
r
t
e
r
i
a
l
p
r
e
s
s
u
r
e
i
s
l
o
w
e
r
t
h
a
n
a
o
r
t
i
c
p
r
e
s
s
u
r
e
i
n
t
h
e
c
o
l
l
167
a
t
e
r
a
l
i
s
e
d
s
e
g
m
e
n
t
)
[
2
1
,
3
2
]
.
1
.
4
E
x
e
r
c
i
s
e
P
h
y
s
i
o
l
o
g
y
E
x
e
r
c
i
s
e
p
h
y
s
i
o
l
o
g
y
i
s
a
b
r
a
n
c
h
o
f
p
h
y
s
i
o
l
o
g
y
,
w
h
i
c
h
s
t
u
d
i
e
s
h
o
w
e
x
e
r
c
i
s
e
a
l
t
e
r
s
t
h
e
s
t
r
u
c
t
u
r
e
a
n
d
f
u
n
c
t
i
o
n
o
f
t
h
e
b
o
d
y
.
E
x
e
r
c
i
s
e
i
s
n
o
w
169
u
s
e
d
a
s
u
r
i
e
s
o
r
a
t
h
e
r
a
p
y
d
u
r
i
n
g
r
e
h
a
b
i
l
i
t
a
t
i
o
n
f
r
o
m
v
a
r
i
o
u
s
i
n
j
i
l
l
n
e
s
s
a
n
d
i
t
i
s
u
s
e
d
a
s
a
p
r
e
v
e
n
t
i
v
e
s
t
r
a
t
e
g
y
t
o
d
e
l
a
y
t
h
e
o
n
s
e
t
a
n
d
p
r
o
g
r
e
s
s
i
o
n
o
f
a
t
h
e
r
o
s
c
l
e
r
o
t
i
c
c
a
r
d
i
o
v
a
s
c
u
l
a
r
d
i
s
e
a
s
e
[
3
3
3
5
]
.
E
x
e
r
c
i
s
e
t
e
s
t
i
n
g
u
s
i
n
g
171
a
t
r
e
a
d
m
i
l
l
o
r
b
i
c
y
c
l
e
c
a
r
d
i
o
l
o
g
y
d
e
p
a
r
t
m
e
n
t
e
r
g
o
m
e
t
e
r
t
o
i
s
t
h
e
c
o
m
m
o
n
l
y
u
s
e
d
i
n
t
h
e
a
s
s
e
s
s
e
x
e
r
c
i
s
e
t
o
l
e
r
a
n
c
e
a
n
d
d
i
a
g
n
o
s
i
s
o
f
i
s
c
h
a
e
m
i
c
h
e
a
r
t
d
i
s
e
a
s
e
.
H
o
w
e
v
e
r
,
e
x
e
r
c
i
s
e
t
o
l
e
r
a
n
c
e
a
n
d
c
a
p
a
c
i
t
y
d
e
p
e
n
d
s
o
n
m
u
l
t
i
p
l
e
f
a
c
t
o
r
s
173
s
u
c
h
a
s
a
g
e
,
s
e
x
,
p
h
y
s
i
c
a
l
/
m
e
n
t
a
l
c
o
n
d
i
t
i
o
n
i
n
g
,
t
i
o
n
s
,
d
i
s
e
a
s
e
s
t
a
t
u
s
,
e
t
c
.
[
3
3
3
5
]
.
D
uri
ng
exe
rci
se,
mu
ltip
le
ph
ysi
olo
gic
m al
e cha
d ng
i es
c are
a enc
ou
nte
red
(Ta
ble
1.7
).
In
ge
ner
al,
car
dia
c
out
put
(C
O)
inc
rea
se
is
du
e
to
a
lar
ger
inc
rea
se
in
hea
rt
rat
e
(H
R)
an
d a
sm
all
er
inc
rea
se
in
str
ok
e
vol
um
e
(S
V).
Du
rin
g
exe
rci
se,
the
CO
ma
y
inc
rea
se
to
a
ma
xi
mu
m
val
ue
of
35
L/
mi
n
(ba
seli
ne
5
L/
mi
n)
[14
].
Mo
st
of
the
inc
rea
sed
car
dia
c
out
put
go
es
to
the
exe
rci
sin
g
175
mu
scl
e
an
d
par
t of
it
go
es
to
the
ski
n
(to
dis
sip
ate
hea
t)
an
d
hea
rt
[14
].
Th
e
inc
rea
se
in
flo
w
to
the
se
org
ans
is
du
e
to
vas
odi
lat
ati
on
an
d
the
flo
w
to
the
gas
troi
nte
sti
nal
org
ans
an
d
kid
ne
ys
dec
rea
ses
(se
co
nd
ary
to
inc
rea
sed
sy
mp
ath
eti
c
act
ivit
y)
[36
39]
.
Th
e
tot
al
per
iph
era
l
res
ista
nce
to
blo
od
flo
w
is
red
uce
d
du
e
to
the
art
eri
ola
r
vas
odi
lat
ati
on
in
the
ske
let
al
mu
scl
e,
ski
n,
an
d
car
dia
c
mu
scl
e.
Th
e
net
res
ult
is a
dec
rea
se
in
tot
al
per
iph
era
l
res
ista
nce
(T
PR
)
[14
].
In
ad
diti
on,
the
re
is
inc
rea
sed
ve
no
us
ret
urn
bec
aus
e
of
inc
rea
sed
mu
scu
lar
act
ivit
y,
wh
ich
in
tur
n
fur
the
r
inc
rea
ses
the
car
dia
c
out
put
.
Th
e
car
dio
vas
cul
ar
an
d
mu
177
scu
lar
cha
ng
es
in
blo
od
flo
w
wit
h
exe
rci
se
is
als
o
co
ntr
oll
ed
by
var
iou
s
fac
tor
s
inc
lud
ing
but
not
lim
ite
d
to
(a)
exe
rci
se
cen
tre
s
in
the
bra
in,
(b)
loc
al
che
mi
cal
cha
ng
es
in
the
mu
scl
e,
(c)
me
cha
nor
ece
pto
rs
an
d
che
mo
rec
ept
ors
in
mu
scl
e,
an
d
(d)
art
eri
al
bar
ore
cep
tor
s
[14
]
(Fi
g.
1.1
2).
1
.
4
.
1
G
e
n
d
e
r
a
n
d
E
x
e
r
c
i
s
e
P
e
r
f
o
r
m
a
n
c
e
Ge
nd
er
dif
fer
en
ces
rel
ati
ng
to
the
car
dio
vas
cul
ar
sys
te
m
in
ter
ms
of
bo
dy
co
mp
osi
tio
n,
me
tab
oli
sm
,
mu
scl
e
mo
rph
olo
gy
an
d
en
do
cri
ne
res
po
nse
s
are
we
ll
rep
ort
ed
an
d
do
cu
me
nte
d
in
the
me
dic
al
179
lite
rat
ure
[40
].
Wo
me
n
ha
ve
sm
all
er
lun
g
vol
um
e
an
d
pul
mo
nar
y
ca
pill
ary
vol
um
e
tha
n
me
n,
res
ulti
ng
in
lo
we
r
ma
xi
mu
m
pul
mo
nar
y
ve
ntil
ati
on
[40
].
Wo
me
n
als
o
ha
ve
a
sm
all
er
he
art,
wit
h a
lo
we
r
fill
ing
vol
um
e,
lo
we
r
ma
xi
mu
m
str
ok
e
vol
um
e
an
d
lo
we
r
car
dia
c
out
put
[40
].
Wo
me
n
are
rep
ort
ed
to
ha
ve
a
lo
we
r
ha
em
ogl
obi
n
co
nc
ent
rati
on,
ha
em
ato
crit
an
d
tot
al
blo
od
vol
um
e;
the
ref
ore
,
the
y
are
at
a
rel
ati
ve
dis
ad
va
nta
ge
for
tra
ns
por
t of
ox
yg
en
to
ske
let
al
mu
scl
e
dur
ing
ex
erc
ise.
Ho
we
ver
,
ge
nd
er
dif
fer
en
ces
be
co
me
les
s
not
abl
e
wh
en
car
dio
vas
cul
ar
par
am
ete
rs
are
ex
pre
sse
d
rel
ati
ve
to
bo
dy
sur
fac
e
are
181
a
an
d
ma
ss
[40
]
(Ta
ble
1.8
).
R
e
s
e
a
r
c
h
n
d
w
o
m
e
n
r
u
n
n
e
r
s
i
n
ONSET OF EXERCISE
Exercising
Centers
In
Brain
Chemical
changes
locally
Stimulate
chemoreceotors
in the muscles
Mechanorecep
tor stimulation
in muscles
Afferent
input
Medullary
cardiovascular
Center
Arteriolar
dilatation in
the muscle
Sympathetic output to heart, veins and arterioles in abdominal organs and
kidneys increases
Parasympathetic output to heart decreases
o
n
m
e
n
a
d
i
c
a
t
e
s
t
h
a
t
w
o
m
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n
t
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n
d
t
o
h
a
v
e
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m
a
l
l
e
r
a
m
o
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n
t
s
o
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l
o
w
t
w
i
t
c
h
f
i
b
r
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s
t
y
p
e
i
n
t
h
e
g
a
s
t
r
o
c
n
e
m
i
u
s
m
u
s
c
l
e
.
T
h
e
g
r
e
a
t
e
r
l
e
183
a
n
b
o
d
y
m
a
s
s
i
n
m
e
n
i
s
a
m
a
j
o
r
d
e
t
e
r
m
i
n
a
n
t
o
f
g
r
e
a
t
e
r
m
u
s
c
l
e
s
t
r
e
n
g
t
h
[
4
0
]
.
M
a
x
i
m
u
m
o
x
y
g
e
n
c
o
n
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u
m
p
t
i
o
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i
s
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n
a
n
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w
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a
n
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m
t
o
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a
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a
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m
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l
o
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g
e
n
c
o
n
s
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m
p
t
i
o
n
[
4
0
]
.
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o
w
e
v
e
r
,
w
h
e
n
t
h
e
s
e
a
r
e
e
x
p
185
r
e
s
s
e
d
i
n
r
e
l
a
t
i
o
n
t
o
l
e
a
n
b
o
d
y
m
a
s
s
,
t
h
e
r
e
a
p
p
e
a
r
s
t
o
b
e
l
i
t
t
l
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s
i
g
n
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f
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c
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n
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e
t
w
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n
t
h
e
s
e
x
e
s
[
4
0
]
.
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i
n
a
l
l
y
,
t
h
e
r
e
a
r
e
c
o
n
s
i
d
e
r
a
b
l
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e
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d
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c
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n
a
n
d
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m
e
n
.
I
n
w
o
m
e
n
w
h
o
t
r
a
i
n
v
e
187
r
y
a
g
g
r
e
s
s
i
v
e
l
y
o
r
t
o
o
h
a
r
d
,
t
h
e
l
u
t
e
a
l
p
h
a
s
e
o
f
t
h
e
m
e
n
s
t
r
u
a
l
c
y
c
l
e
i
s
s
h
o
r
t
e
n
e
d
a
n
d
e
v
e
n
t
u
a
l
l
y
t
h
e
c
y
c
l
e
i
s
l
o
s
t
(
a
t
h
l
e
t
i
c
a
m
e
n
o
r
r
h
o
e
a
)
[
3
3
,
4
0
]
.
1
.
4
.
2
A
g
e
a
n
d
E
x
e
r
c
i
s
e
P
e
r
f
o
r
m
a
n
c
e
A
g
i
n
g
i
s
a
n
o
r
m
a
l
b
189
i
o
l
o
g
i
c
a
l
e
f
i
n
e
d
p
r
o
c
e
s
s
,
w
h
i
c
h
i
s
i
n
e
v
i
t
a
b
l
e
.
A
g
i
n
g
h
a
s
b
e
e
n
d
a
s
p
r
o
g
r
e
s
s
i
v
e
l
o
s
s
o
f
p
h
y
s
i
o
l
o
g
i
c
a
l
c
a
p
a
c
i
t
i
e
s
t
h
a
t
c
u
l
m
i
n
a
t
e
s
i
n
d
e
a
t
h
[
4
0
4
2
]
.
191
Tab
le
1.8
Gen
der,
car
dio
vas
cula
r
syst
em
and
exe
rcis
e
phy
siol
ogy
[40]
Females have a sm
Females have a lo
Var
iou
s
nat
ura
l
an
d
pat
ho
ph
ysi
olo
gic
al
ch
an
ges
oc
cur
dur
ing
the
co
urs
e
of
life
[41
,
42]
(Ta
ble
1.9
).
Th
e
co
m
mo
nes
t of
193
the
se
are
los
s
of
hei
ght
,
gre
yin
g
hai
r,
ch
an
ges
in
vis
ion
,
red
uct
ion
in
lea
n
bo
dy
ma
ss,
wri
nkl
ing
of
ski
n,
etc
.
[41
,
42]
.
Ma
ny
int
ere
sti
ng
the
ori
es
ha
ve
be
en
im
pli
cat
ed
to
ex
pla
in
the
pro
ces
s
of
agi
ng.
Ho
we
ver
,
ho
w
lon
g
an
ind
ivi
du
al
liv
es
de
pe
nd
s
on
mu
ltip
le
fac
tor
s
inc
lud
ing
her
edi
ty,
en
vir
on
me
nt,
ind
ivi
du
al
atti
tud
e
to
wa
rds
he
alt
h
an
d
my
an
d
ph
ysi
olo
gy
is
ext
ens
ive
life
an
d
fin
all
y
acc
ess
to
he
alt
h
ser
vic
es
[41
,
42]
.
,
fas
cin
ati
ng
an
d
ex
citi
ng.
Ho
we
ver
,
ex
plo
itin
g
the
ir
rol
e
in
cli
nic
al
pra
cti
ce
req
uir
es
a
tho
rou
gh
un
der
sta
ndi
ng
of
bas
1
.
5
C
o
n
c
l
u
s
i
o
n
Ca
rdi
ac
an
ato
195
ic
pri
nci
ple
s,
me
ch
ani
sm
s
an
d
fun
cti
on
alit
y.
Ap
pre
cia
tin
g
an
d
un
der
sta
ndi
ng
the
an
ato
my
an
d
ph
ysi
olo
gy
an
d
rec
og
nis
ing
the
ir
ch
an
ges
wil
l
hel
p
not
onl
y
in
ant
ici
pat
ion
of
dis
eas
e
but
als
o
in
ap
pre
cia
tin
g
the
sig
ns
an
d
sy
mp
to
ms
pro
du
ce
d
by
the
m.
Maximum
Maximum
Cardiac o
Blood pre
Vascular
Total cho
Triglycer
LDL
HDL
Maximum
Respirato
Vital cap
Expirator
1.
diolstrength
Muscle
ogy
Pulmonary
Fun diffu
dam
Alveolar surface
enta
Chestlswall struc
and
Alveolar elastic
Prac
tice, bloo
Pulmonary
Vol.
Lean1,body mass
2nd tissue
Adipose
edn.
BasalMos
metabolic
bySleep
Yea
r
Cognitive
functi
Boo
k,
R St
e Lou
is,
f
e pp.
3
r
25.
e
Ed
n war
c
ds
e WD
(19
s
84)
Ana
Call tom
aha y of
the
n
JA, Car
Kay diov
ascu
JD
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91) Syst
Fou em;
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