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1915 Law of The Heart Lecture by Starling
1915 Law of The Heart Lecture by Starling
ON THE
BY
ERNEST H. STARLING
M.D., Sc.D. (Cambridge and Dublin), F.R.C.P., F.R.S.
THE LINACRE L EC TU R E
ON
LAW OF T H E HEART
G I V E N A T C A M B R ID G E , 1915
BY
E R N E ST H. STARLING
M .D ., Sc.D. ( C a m b r i d g e a n d D u b l i n ) , F.R.C.P., F.R.S.
W IT H D I A G R A M S
1918
T H E LAW O F T H E H EART.
h a s sh o w n th a t in th e n o rm a l in d iv id u al a t re s t th e left v e n tric le
p u m p s o u t a b o u t 3 litre s o f b lo o d p e r m in u te. D u rin g m o d e ra te
m u sc u la r ex e rc ise th is q u a n tity m a y rise to 12 litres, o r in an
a th le tic in d iv id u a l u n d e r ta k in g v e ry h a rd w o rk to as m u c h as
21 litre s p e r m in u te. T h is la st figure m e a n s th a t th e h e a rt can
p u m p th e to ta l b lo o d c o n ta in e d in th e b o d y ro u n d th e w h o le
c irc u la to ry sy ste m six o r seven tim e s in th e co u rse o f a m in u te ,
a n d th is in sp ite o f th e fact th a t d u rin g m u sc u la r e x e rc ise th e
a rte ria l p re ssu re a g a in st w hich th e h e a rt h as to w o rk is c o n s id e r
a b ly h ig h e r th a n in c o n d itio n s o f rest, b e in g e q u iv a le n t in th e
la tte r ca se to a b o u t n o m m . H g . a n d in th e fo rm er to 150
to 160 m m . H g .
A sim ila r p o w er o f a d a p ta tio n is sh o w n in cases o f d ise ase .
W e can im ita te th e d ise a se d c o n d itio n e x p e r im e n ta lly b y d e
s tro y in g a n a o rtic valve, so th a t th e a n im a l su ffers from a o rtic
re g u rg ita tio n , o r b y d im in ish in g th e lu m en o f th e a o rta b y m e an s
o f a lig a tu re , so a s to p ro d u c e artificial a o rtic ste n o sis. T h e
a v e ra g e a rte ria l p re ssu re u n d e rg o e s n o a lte ra tio n as a re s u lt of
e ith e r o f th e se p ro c e d u re s, a lth o u g h th e le ft v e n tric le h a s to p e r
form tw o o r th re e tim e s as m u c h w o rk as it d id b efo re in o rd e r
to m a in ta in th e b lo o d p re ssu re a t its p rev io u s level. T h e circu
la tio n o f b lo o d th ro u g h th e b o d y re m a in s th e re fo re unaffected.
T h is a d a p ta tio n o f th e h e a r t to v a ria tio n s in th e d e m a n d m a d e
u p o n it o ccu rs e q u a lly w ell a fte r to ta l d e s tru c tio n o f th e n erv e s
c o n n e c tin g th e h e a r t w ith the' c e n tra l n e rv o u s sy stem . So we
m u s t c o n c lu d e th a t th e g o v e rn o r m e ch a n ism , in v irtu e o f w h ic h
th e h e a rt is a b le to d o m o re o r less w o rk , a c c o rd in g to th e
a m o u n t o f b lo o d w hich h a s to be s e n t on a n d th e re sis ta n c e to
th e flow p re s e n te d b y th e a rte ria l p ressu re, m u s t b e s itu a te d in
th e w alls o f th e h e a r t its e lf an d p re su m a b ly in th e m u sc le fib res
o f w h ich th e s e a re com p o sed . I t is th e n a tu re o f th is m e c h a n ism
w h ich I w ish to d iscu ss w ith y o u to-d ay .
I n o rd e r to ' in v e stig a te it w e m u s t b e a b le in th e iso la te d
h e a r t to c o n tro l e x a c tly all th e m e c h a n ic a l c o n d itio n s o f th e
h e a r t, e.g. volu m e, p ressu re, a n d te m p e ra tu re o f e n te rin g b lo o d ,
a n d th e re sis ta n c e to th e outflow offered b y th e a rte ria l re sist
6 TH E L IN A C R E L E C T U R E
th a t th e re is n o d a m m in g u p o f th e b lo o d o n th e v en o u s sid e
n o r failure o f th e h e a rt to send th e b lo o d o n from th e v en o u s to
th e a rte ria l side. In th is case also th e r e m a y b e so m e in c re a se
o f c o ro n a ry flow, w h ich is n o t h o w ev e r so m a rk e d as w h en
w e th ro w m o re w o rk o n th e h e a r t b y in c re asin g th e a rte ria l
resistan ce.
D o e s th is a u to m a tic a d a p ta tio n o f th e w o rk o f th e h e a r t to
th e d e m a n d s m a d e u p o n th is o rg a n m e a n th a t a g r e a te r p ro p o r
tio n o f th e chem ical en e rg y o f th e c o n tra c tin g m u scle is tr a n s
form ed in to m ech an ical en e rg y , o r d o es th e h e a r t m u sc le a c tu a lly
se t free m o re e n e rg y w h en its w o rk is in c re a se d ? T h is q u e stio n
h as b een an sw e re d b y L o v a tt E v an s. J u s t as th e r e s p ira to ry e x
c h a n g e s o f th e w hole b ody, viz. th e in ta k e o f o x y g e n a n d th e
o u tp u t o f ca rb o n ic acid, m a y b e u se d as a m e a su re o f th e ch em ical
ch an g es, a n d o f th e to ta l e n e rg y se t free in th e b o d y , so th e re
s p ira to ry e x c h a n g e s o f th e h e a r t m a y be u tilize d for d e te rm in in g
th e to ta l e n e rg y se t free in th is o rgan. In th e h e a rt-lu n g p re p a ra
tio n th e re s p ira to ry m e ta b o lism o f th e h e a rt c a n b e in v e stig a te d b y
m e a su rin g th e o x y g e n ta k e n u p from th e lungs, a n d th e c a rb o n ic
acid g iv e n o u t b y th e lu n g s. W h e n w e d o th is w e find th a t a n y
in c re ase in th e w o rk o f th e h e a rt, h o w ev e r cau sed , w h e th e r b y a
rise in th e a rte ria l p re ssu re o r b y a m o re ra p id v en o u s inflow, is
a tte n d e d b y a c o rre sp o n d in g a u g m e n ta tio n o f th e to ta l ch em ical
changes. T h is is sh o w n in th e follow ing ta b le s :
EXP. I . A R T E R IA L R E S IS T A N C E V A R IE D . IN F L O W C O N S T A N T .
H eart, 57 gm s.
130 8 371 60
* I have recently found that in these experiments both the work done and the
mechanical efficiency were underestimated. In exp. 2 (p. 11) the mechanical effi
ciency w ith an inflow of 62 litres per hour was probably about 25 per cent, and the
other figures must be increased in proportion.
T H E L A W O F TH E H E A R T 11
EXP. 2. A R T E R IA L R E S IS T A N C E C O N S T A N T . V E N O U S IN F L O W
V A R IE D . , .
H e a rt, 53 g m .
62 331 + 86 9 70 60
In these tables the oxygen consumption serves as an index o f the total chemical
changes and o f the chemical energy set free. T his may be measured in calories,
and compared w ith the total work done by the heart in the same time, the result
being the mechanical efficiency o f the heart as a machine.
The work o f the heart in kilogrammetres was determined by m ultiplying the
output per hour by the average arterial pressure (in metres o f blood).
In the first experiment the work o f the heart was varied by altering the arterial
resistance, in the second by altering the venous inflow.
A.P. = arterial pressure. V.P. = venous pressure.
The heart volume is measured from an arbitrary base line, viz. the volume at
the beginning o f the experiment. Note the marked enlargement o f the heart which
attends increase in venous inflow.
W e th u s find th a t th e h e a r t h a s a m a rv e llo u s p o w e r o f a d
ju s tin g n o t o n ly its o u tp u t o f m e ch a n ica l e n e rg y b u t also its to ta l
e n e rg y c h a n g e s to th e w o rk w h ich is im p o sed u p o n it b y th e
m e ch a n ica l c o n d itio n s o f th e c irc u la tio n . In m u sc u la r ex e rc ise
fo r in sta n c e , w h en th e inflow o f b lo o d in to th e la rg e v ein s is
in c re a se d b y th e p u m p in g a c tio n o f th e m uscles a n d th e a rte ria l
p re ssu re is raise d b y th e c o n tra c tio n o f th e sm all a rte rie s g o in g to
th e a lim e n ta ry ca n al, th e h e a rt picks u p all th e b lo o d w hich co m es
to it a n d se n d s it o n w ith in c re a se d force in to th e a rte ria l sy stem ,
w h en c e it ca n be d is trib u te d in ric h m e a su re to th e c o n tra c tin g
m u sc le s ; so th a t, as K ro g h has show n, its o u tp u t p e r m in u te m a y
rise from 3 litre s to th e e n o rm o u s fig u re o f 21 litres. W e m ay
form so m e id e a o f th e p ro cesses in v o lv ed in th is a d a p ta tio n b y
c o m p a rin g th e h e a r t to a m o to r cycle. H e r e also th e e n e rg y is
d e riv e d from th e o x id a tio n o f c e rta in co m b u stib le su b stan ce s, in
th is case p e tro l sp irit. W h e n o n ce a d ju ste d th e m o to r cy c le w ill
12 TH E L IN A C R E L E C T U R E
Fm. 3. Effect o f alteration in venous supply on volume o f heart. (Read from left
to right.)
Dog, 5*15 kilos; heart, 67 gms.
O .P. per
O .P. per beat
A.R. B.P. V .P. Rate. O .P . beat calc. observed.
A. 100 124 95 22 86 3'9 57
B. 100 130 145 22 140 6-4 8-0
C. 100 122 55 22 33 1-5 2-5
N ow h e re a re tw o co n d itio n s in w hich th e w o rk o f th e h e a rt
is increased a n d in w h ich th is o rg a n a d a p ts its e lf b y in c re a sin g
th e chem ical c h a n g e s in its m u scle a t ea c h c o n tra c tio n to th e in
cre ase d d e m a n d s m a d e u p o n it. I t is e v id e n t th a t th e re is o n e
fa c to r w hich is co m m o n to b o th cases, a n d th a t is th e in c re ase d
i8 TH E L IN A C R E L E C T U R E
v o lu m e o f th e h e a rt w hen it b eg in s to c o n tra c t. S o th a t w e m a y
m ak e th e follow ing g e n e ra l sta te m e n t. W it h in p h y s io lo g ic a l
lim its th e la r g e r th e v o lu m e o f th e h ea rt, th e g r e a t e r a re th e
e n e r g y o f it s co n tr a c tio n an d t h e a m o u n t o f c h e m ic a l c h a n g e
a t ea c h co n tra c tio n .
40
F i g . 6. Diagram showing relation between fillin g , in itia l tension, and final tension
in a heart contracting isom etrically (abscissa = volume o f heart contents,
[T"_ ordinates = intracardiac pressure in mm. Hg).
h e a r t is in c re a se d th e re is a c o rre sp o n d in g rise in th e h e ig h t
of th e p ressu re d e v e lo p e d d u rin g c o n tra c tio n . I t w as fo rm e rly
t h o u g h t o n th e b asis o f F r a n k s e x p e rim e n ts th a t th e e sse n tia l
fa c to r d e te rm in in g th e s tre n g th o f c o n tra c tio n o f th e card iac m u scle
-was th e te n sio n it w as u n d e r a t th e b e g in n in g o f th e c o n tra c tio n .
I t w ill be o b se rv e d , how ever, from th is d ia g ra m th a t th e rise o f
diastolic te n sio n is a t first n eg lig ib le a n d th a t as soon as th e
p re s s u re w ith in th e v e n tric le d u rin g d ia sto le b e g in s to in c re a se
thus serves to measure the amount o f tension produced in the heart muscle at each
contraction. The arrangement o f the experiment is shown in the diagram (fig. 4)
a nd the actual record o f a series o f contractions w ith different fillin g is given in fig. 5.
20 THE L INACRE LECTU RE
a p p re c ia b ly , so as to ca u se a passiv e s tre tc h in g o f th e m u s c le
fibres, th e a b s o lu te te n sio n d e v e lo p e d d u rin g c o n tra c tio n rise s
v ery little m o re o r m a y even fall. T h e r e ca n b e n o q u e s tio n
th erefo re th a t it is th e v o lu m e o f th e v e n tric le s a t th e b e g in n in g
o f co n tra c tio n ra th e r th a n th e p re ssu re w ith in th e ir c a v itie s,
w hich d e te rm in e s th e a m o u n t o f e n e rg y se t free d u rin g th e c o n
tractio n . T h is free en e rg y d isp la y s its e lf as c o n tra c tile stress*
a n d causes a rise o f p ressu re w ith in th e c a v ity o f th e v en tricle.
Pressure
30 60 90 120 150 180 210 240 270 ^ , H # .
* ------------- ------------ '--------------------------- 0
<
o 30-
c
3n>
F i g . 7 . See
text. The abscissae represent pressure in m m . Hg, and the ordinates
measured from above the capacity of the ventricles. The left-hand curve re
presents the condition of the heart during diastole. The right-hand curve
that during systole.
i
fO
o
>
u.
0
T
>-
1ca
3
o
3
Ci
a.
F ig .
8. Curve showing the relation o f tension, in itia l length, and energy evolved
in an isometric tw itch. (Note the resemblance o f this curve to that o f the
heart in fig. 6.)
A , Condition o f maximum shortening.
B, Condition o f maximum extension of muscle in its normal relationships in the body.
P R IN T E D IN GR EAT B R IT A IN BY T H E U N IV E R S IT Y P R E S S , A B ER D EEN .