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Vendrely, R., Knobloch-Mazen, A., and Vendrely, 0., in The Cell Nucleus,

CHEMISTRY OF MUSCLE CONTRACTION

ADENOSINE TRIPHOSPHATE AND PHOSPHORYLCREATINE AS ENERGY SUPPLIES FOR
SINGLE CONTRACTIONS OF WORKING MUSCLE
By DR. D. F. CAIN, A. A. INFANTE* and PROF. R. E. DAVIES
Departments of Animal Biology and Molecular Biology, School of Veterinary Medicine,
University of Pennsylvania, Philadelphia

acids9’10 the phosphorus in the total acid-insoluble residue of

C ONTRACTING muscle is one of the most fascinating
‘machines’ in Nature, but, despite an enormous effort, it has
so far not been possible to find the immediate source of energy
muscle measured either as inorganic phosphate, phosphatido-
peptide phosphorus, phospholipid phosphorus, ribonucleic or
which allows this mechano-chemical system to turn chemical deoxyribonucleic acids11. Other relevant experiments12 have
free energy into external work. Following the work of Fletcher been reviewed by Davies and Cain13.
and Hopkins1 in 1907, Parnas and Wagner2 in 1914, Meyerhof3 Since we had found that inorganic phosphate was certainly
in 1920, it was generally accepted that the energy for muscle liberated during a single muscle contraction (Table 1), and no
contraction came from the breakdown of glycogen to lactic acid. other compound could be observed to account for it, we re-
However, in 1930, Lundsgaard4 showed that iodoacetate-treated investigated the possibility of a breakdown of PCr using gentler
muscles could contract many times without producing lactic methods of extraction and more precise methods of analysis.
acid. During these contractions, phosphoryl- creatine (PCr) was These experiments showed that PCr in fact liberates creatine and
found to break down and thus appeared to be the source of inorganic phosphate during a single contraction9’10’13'14 (Table
energy. Lohmann6, in 1934, found that the eo-factor for the 1). This finding has now been confirmed by Fleckenstein16 and
breakdown of PCr in dialysed cell-free muscle extracts was Mommaerts16. In normal rectus abdominis muscles of the female
adenosine diphosphate (ADP). Thus, it was generally accepted frog the inorganic phosphate and creatine content are
that the dephosphorylation of adenosine triphosphate (ATP) to sufficiently low (that is, 2-5 and 5-0 pmole/g) that changes can
produce ADP and inorganic phosphate was the initial reaction be observed during contraction, hut the concentration of PCr is
for the provision of energy for contraction in spite of the fact so high it is not possible to observe the expected changes with
that the content of ATP remained unchanged except in extreme sufficient certainty. However, if the concentration of PCr is first
fatigue. This view was supported by experiments with isolated lowered from about 12 to about 2-5 pmole/g by treatment with
enzymes, muscle protein solutions, fibrils made from 2,4-dinitrophenol (DNP), (under conditions where the muscle
reconstituted actomyosin, and glycerinated musele models can still do several complete contraction cycles) then the
which strongly suggest that ATP is in fact the immediate source expected change in PCr can
of energy for muscle contraction6.
However, in 1954, Fleckenstein, Janke, Davies and Krebs’
published experiments showing that frog rectus abdominis
muscles could contract and do work at 0° C without measurable
net change of energetically equivalent quantities of ATP, ADP
or PCr (that is, about 0-5 pmole/g at 50 per cent efficiency),
while Mommaerts8 reached similar con* Predoctoral Fellow of the
IT.S. National Institutes of Health.
elusions on the basis of experiments with turtle leg muscles at 0°
C in which measurements were made of ADP, adenosine
monophosphate (AMP), creatine, creatinine and pyruvate during
contraction and relaxation. Since then, many other phosphorus
compounds have been tested and found not to undergo sufficient
net changes during contraction. These include carnosine di- and
mono-phosphates9, all the inosine, guanosine and cytosine
phosphates, phos- phoenolpyruvate, all the phosphoglyceric

© 1 962 Nature Publishing Group

 No. 4851 October 20, 1962 NATURE 215
Table 1. CHANGES IN INORGANIC PHOSPHATE (Pi) AND FREE CREATINE (CT)IN NORMAL FROG RECTUS ABDOMINIS MUSCLES FOLLOWING ELECTRICAL
STIMULATION AT 0° C
Average external stimulation (si
Pairs of AV\ ACT work (12 supramaxii
Type of exp. muscles (fimolejg muscle) (/umole/g muscle) (g-cm/g muscle) pulses/sec)
Control-vs-activation Control-vs- 13 + 0-08 ±0*11 + 0*11 + 0*14 0-10 <0-2
shortening Control-vs-contraction 9 -0-08 ±0-20 + 0*02 + 0*21 0 0*5 -1-0
Control-vs-contraction + relaxation + contraction 21 + 0-64 ±0-14 + 0*83 + 0*20 120 10-2*5
Control-vs-control 10 + 1*20 ±0-17 + 1-22 + 0-30 210 4*0-6*0
Contraction-vs-contraction + relaxation 12 0-09 + 0*10 0-29 + 0*26 0 0*0
——
7 + 0*08 ±0-14 120* 1-0-2*5
* Both muscles of the pairs did the same amount of wort.
The paired muscles from female Rana pipit,ns were dissected free, separated, and incubated in physiological bicarbonate saline solution at 25° C for 1-2
h. They were cooled to 0° C for 5 min and used as follows: In the activation experiments the muscles at rest length were stimulated and then frozen in liquid
'Freon' (CF,C1, + CF.Cl) at —172° C as shortening began but before much external work had been done. In the shortening experiments the muscles were
stimulated to shorten to one-half of their rest length under no external load and then frozen. Control- control values are for experiments in which neither
member of the paired muscles was stimulated. Changes in Pi brought about by relaxation were studied by freezing one muscle at the eight of 'after-loaded'
contraction and permitting its pair to contract and then relax under load before being frozen. Muscles were extracted in aqueous methanol at — 35° C for 7 days
and the Pi and Cr measured by the methods of Wahler and 'Wollenberger n, and Eggleton, Elsden and Gough18. Here and elsewhere, except when noted, all
assays were done in duplicate. Results
± standard error of means.

Table 2. EFFECT OF ELECTRICAL STIMULATION IN HYPERTONIC very clear correlation with breakdown of PCr was found (Table
SOLUTION AT 0°C ON THE PHOSPHORYLOREATINE (PCr) LEVEL OF
FROG RECTUS ABDOMINIS MUSCLES 3). Our direct chemical measurements show that in this muscle
No. of pulses JPCr/pulse the breakdown of PCr can he associated with the work done and
Pairs of muscles (at 12/sec) (/umole/g muscle) is rather independent of the amount of shortening that occurs.
4 12 0-00 + 0-04
5 24 + 0*01 + 0-01 It is of interest that DNP-treated muscles, which have low
5 36 0-00 + 0*01 PCr but high creatine and inorganic phosphate, work with a
These muscles were treated with 0-25 mM DNP at 20° C for 25 min reduced stoiehioehemieal efficiency (work per mole) (Tables 1
and then for 20 min at 0° C in DNP-bicarbonate saline made
hypertonic with 440 mM sucrose. Other conditions were as in Fig. 1. and 4, Fig. 1) which might well be expected on thermodynamic
The work performed by these muscles was less than 3 g-cm/g muscle grounds.
Table 3. ASSOCIATION OF PHOSPHORYLOREATINE (PCr) BREAKDOWN WITH In a mechano-chemical system the relation between output
EXTERNAL WORK BUT NOT WITH DEGREE OF SHORTENING Per cent
shortening External Stimulation of heat and work done depends on the entropy and free energy
It “ IB zdPCr work time (sec) of the chemical reactions, the efficiency of energy transduction
Pairs of ----------X 100*
lo
(mnole/g (g-cm/g (12 pulses/ and the absolute temperature. As Wilkie21 has pointed out, when
muscles muscle) muscle) sec)
Constant shortening—varying external work work is done by ATP usage under standard conditions at about
10 19*6 ± 2*24 -0-23 + 0-15 21*7 + 1-9 0-6 ±0-1 38 per cent efficiency, the reaction is thermally neutral. This
10
10
20-2 + 2*82 -0*36 + 0-16
20-5 + 2*27 -0*47 + 0*26
30*8 + 2*3
46*6 + 2*3
1*2 ±0*1
1*6 ±0*2
type of situation may account for Hill’s observation22 of the
8 21*1+1*28 -0*58 + 0-18 55*2 + 0*9 2-0 ± 0*4 striking appearance of work without extra initial heat production
5 19*7 + 3*13 -0-77 + 0*24 64*2 + 6*0 2-6 ±0*5 (above shortening heat) during the rising phase of a contraction,
Varying shortening—approx, constant external work the time when we observe a breakdown of PCr. Assuming that
11 10*7 + 0*81 -0*44 + 0*22 36*3 + 4*7 1*9 ±0-4
11 18*5 + 0*75 -0-44 + 0*16 37*3 + 4*6 1-0 ±0*2 results on sartorius muscles apply to rectus abdominis, our
11 23*4 + 0*31 -0*48 + 0*21 39-3 + 5-1 1*2+ 0*2 findings also show that the Fenn effect is to be expected; that is,
9 29*5 + 1*08 -0*54 + 011 50-8 + 4-3 2-0 ±0-4
*la is rest length ; ls is length after shortening.
“ . . . when work is
By varying the load, the muscles were made to do either different
quantities of work while shortening the same amount, or the same amount of
work while shortening to different lengths. The amounts of shortening and
external work performed were determined from kymograph tracings. The
muscles were treated as described in Fig. 1.

be observed to occur during a single contraction. The

experimental techniques have been described previously9’10 A*.
The results are shown in Tables 1-3 and Fig. 1. It is clear that
muscles made to do varying amounts of work broke down
varying amounts of PCr to inorganic phosphate and creatine
with a linear relationship between the amount of work and the
amount of breakdown (Table 1, Fig. 1). Relaxation was not
associated with any measurable change (Table 1). It is of interest
that the observed breakdown of PCr is associated very much
more with work than with either activation or shortening. The
breakdown on activating the muscle and rapidly freezing it
before it was allowed to do any work was quite small (Table 1).
This was confirmed by stimulating the muscle in hypertonic
saline (Table 2), where muscles are still activated but do not
contract20. Similar small changes occurred in muscles which
shorten against zero external load (Table 1). Furthermore, Fig. 1. Relationship of phosphoryloreatine breakdown to the amount of
muscles performing an approximately constant amount of work external work performed by ‘after-loaded’ frog rectus abdominis muscles
contracting once or twice against a constant load to different degrees and for
with large changes in the degree of shortening showed no different times at 0° C. These muscles were isolated and incubated as
significant change in PCr breakdown associated with the described in Table 1, then treated with 0-25 mM DNP in bicarbonate saline
solution for 40 min at 20° C and cooled to 0' C for 5 min. PCr was converted
shortening (Table 3), whereas for constant shortening and to creatinine by molybdate-catalysed acid hydrolysis and measured
varying work a according to Barker and Ennor1*. The figures in parentheses refer to the
numbers of pairs of muscles used. Means ±S.E.
Slope of line: 2,600 calories of external work/mole PCr

© 1 962 Nature Publishing Group

 216 NATURE October 20, 1962 VOL. 196

Table 4. PRODUCTION OF INORGANIC PHOSPHATE (Pi) WITHOUT CHANGE IN phoryltransferase could be inhibited in situ and thus allow a
PHOSFHORYLCKEATINE (PCr) AFTER THREE SMALL CONTRACTIONS AT 0° IN
FROG RECTUS ABDOMINIS MUSCLES PRETREATED WITH DINITROPHENOL direct demonstration of the primary role of ATP in muscular
(DNP) PLUS FLUORODINITROBENZENE (FDNB) contraction. Kuby and Mahowald28 had found that crystalline
External
Pairs work creatine phosphoryltrans- ferase is completely inhibited by l-
(g-cm/g APCr
of muscle) (pmole/g (/amole/g fluoro-2,4-dini- trohenzene (FDNB), and we have now used
Type of exp. muscles muscle) muscle) this inhibitor to find the changes in ATP during single
Control-vs-
contractions 27 81 ±5 — 010 + 0-08 .—-— contractions which have been sought so often during the past
Control-vs- thirty years29. When isolated muscles were pre-treated with
contractions 12 79 ±8 -0*17 + 0-08 + 1-23 ±0*48 FDNB, the number of normal contractions which could he
—.—•
Control-vs-control 4 0 0*17 + 0*17
Control-vs- performed was reduced from more than 30 to about 3, yet even
contractions 4 76±8 -1*02 + 0*28 •— — when they would no longer contract, they still contained the
(DNP but no FI)Nil)
normal quantities of PCr found in untreated muscle.
The paired muscles were isolated and incubated as in Table 1. They were Table 4 shows that after treatment with DNP + FDNB,
then placed in bicarbonate saline solution containing 0-25 mM DNP for 30
min at 20° C followed by 20 min at 0° C in saline containing both 0-25 mM inorganic phosphate increased on contraction although PCr
1)N11 and 0-38 mM FDNB. These salines were gassed with 5 per cent CO, + remained unchanged. Table 5 gives results with FDNB alone
95 per cent If,. Total time of stimulation 4-6 sec.
which show that ATP was split, hut that the net amount of ADP
produced was much less than the net ATP breakdown which
Table 5. BREAKDOWN OF ADENOSINE TRIPHOSPHATE (ATP) TO FORM
ADENOSINE DIPHOSPHATE (ADP) AND ADENOSINE MONOPHOSPHATE itself was less than the inorganic phosphate production
(AMP) DURING CONTRACTION OF FROG RECTUS ABDOMINIS MUSCLES observed in the earlier experiments given in Table 1.
AFTER TREATMENT WITH FLUORODINITROBENZENE (FDNB)
In these earlier experiments with normal muscles, the
ADP AMP
(pmoles/g (pmoles/g (pmoles/g Lohmann reaction6 was apparently the over-all energy-
muscle) muscle) muscle) supplying mechanism; that is: actomyosin ATPase
Single contraction ATP -------------------------- >ADP+inorganic phosphate + work, and
Control 1-25 0-64 0-10 creatine phosphoryltransferase PCr+ADP — —
After 1 contraction 0-81 — ---------------- —*ATP + creatine
Change + S.E. for 9 0-90 0-24
pairs -0-44 ±0-046
Double contraction + 0-26 ±0-023 + 0-14 + 0-027 Thus, the changes in inorganic phosphate and creatine
Control 1-24 should be identical as is shown in Table 1. The apparent
After 2 contractions 0-61 0-07
Change ± S.E. for 3 0-59
0-88
discrepancy shown in Table 5 can he resolved by consideration
pairs 0-41
— 0*65 ±0-045 of the action of myokinase. This enzyme catalyses tho reaction:
+ 0-27 ±0-051
External work ^ 100 g-cm/g muscle/contraction. + 0-34 ±0-037 myokinase
2 ADP ------------------------------ s- ATP + AMP
The muscles were Isolated and incubated as described in Table 1 then
treated with 0-38 mM FDNB for 40 min at 0° C. ATP, ADP and AMP were
measured fluorometrically according30 to a modification of the method
described by Estabrook and Maitra . AMP results are based on single
analyses. In some experiments, the ATP contents of control and contracted
muscles were also measured by the use of the specific firefly luminescence
technique with a photomultiplier coupled to a recorder81.
done by the muscle, an extra supply of energy is liberated . . .
which is roughly proportional to the work . . . ”23. Thus there Thus, the combined action of actomyosin ATPase and
will be an extra heat output after tho shortening while, inter alia, myokinase leads to the generalized equation given in Table 6.
the PCr used by working muscle is being re-synthesized. This generalized equation fits the experimental results
However, the existence of shortening heat (apart from work remarkably well for both conditions tested. These findings,
heat) has recently been questioned21 so the Ferm23-26 effect together with those we have obtained previously, demonstrate
might be directly related to the breakdown of PCr rather than to clearly and directly that the energy is delivered from a break-
its re-synthesis. down of ATP. They also make it clear that, in normal working
Lundsgaard4 showed, many years ago, that the energy for muscle, creatine phosphoryltransferase makes available the
muscular work in a series of contractions could be directly energy stored in phosphorylcreatine, but that in extreme
related to the splitting of PCr, and this has recently been conditions myokinase can also be observed to operate in muscle.
confirmed for large series of twitches24-26. However, it was not Myokinase activity is important because by cyclic
possible to determine whether the breakdown occurred during regeneration of ATP from ADP free energy is in effect released
contraction or relaxation or whether the energy was delivered at from the breakdown of both the labile phosphates of some of the
the beginning or throughout the contraction cycle, whereas our ATP. The enzyme also maintains low levels of ADP, which is an
experiments show that this chemical change occurs while the inhibitor of myosin ATPase32, and it keeps the ratio of ATP to
work is being done by tho muscle. It is clear, and has been ADP high. On thermodynamic grounds, this is advantageous for
pointed out by Hill27 and others, that the splitting of PCr is not free energy liberation from ATP splitting.
proof of the breakdown of ATP. However, this breakdown Our results fit the sliding-filament theory33. If the tension-
should be observable if creatine phos- generating system depends on the formation
Table 6. STOICHIOMETRY OF THE CHEMICAL REACTIONS WHICH OCCUR DURING CONTRACTION AT 0° C OF FROG RECTUS ABDOMINIS MUSCLES
TREATED WITH FLUORODINITROBENZENE
Generalized equation for the combined action of actomyosin ATPase and myokinase n ATP
-------------------------- >- (n~m) ADP + m AMP + (n + m) Pi.
One contraction of frog rectus abdominis (All quantities in /imoies/g muscle)
(0-44 ±0-046) ATP --------------- -K0-26 ± 0-023) ADP + (0-14 ± 0-027) AMP + (0-64 ± 0-14) Pi *
On the basis of the figures for ATP and ADP, the values calculated from the generalized equation are 0-18 AMP and 0-62 Pi.
Two contractions of frog rectus abdominis (All quantities in pmoles/g muscle)
(0-65 + 0-045) ATP ---------------- ► (0-27 + 0-051) ADP + (0-34 ± 0-037) AMP + (1-20 ± 0-17) Pi*
The calculated values are 0-38 AMP and 1-03 Pi.
* Since we have not so far measured changes in ATP, ADP, AMP, PCr, Cr, and Pi all on the same muscle pairs, the changes in Pi are taken from the
results on normal muscle given in Table 1.

© 1 962 Nature Publishing Group

 No. 4851 October 20, 1962 NATURE
of cross-links between the side branches of the myosin and the
actin, then fewer links would be formed during rapid unloaded
contractions and more would be formed during slow, working
contractions. Thus different amounts of energy would be
required. The demonstration that myokinase is operating during
the actual contraction means that the same ATP can be used
several times. Such a re-use of a small amount of ATP is
consistent with the view that the side-branches must form bonds,
develop tension, and break the bonds many times during a single
contraction by a mechanism involving ATP splitting; it is
inconsistent with the view that a given amount of ATP binds on
to the muscle protein and causes a complete contraction by an
entropie electrostatic mechanism84. In this case, the ATP would
remain bound during the whole contraction and would only be
released at the end as ADP + inorganic phosphate. Thus, neither
ATP splitting nor myokinase action would occur during the
contraction.
Our results strongly support the view that the shortening of
muscle is due to the summation of effects caused by a small
number of bonds which are formed and broken repeatedly. It
follows that an ATP-dependent series of micro-entropic
contractions of the myosin side pieces after they had linked
asymmetrically with the actin might well account for the overall
process of muscle contraction by the sliding filament
mechanism.
In summary, these results show that, with frog rectus
abdominis muscles when work is done, ATP is used during
single contractions, and that this ATP is normally rapidly
reconstituted by the action of creatine phosphoryltransferase and
myokinase while the contraction is taking place. There is a linear
relationship between the external work done and the cleavage of
high-energy phosphate compounds, but very little energy from
this source is required for activation and shortening per se in
these muscles.
We thank Miss Dzintra Klaupiks, Messrs. W. J. Brunton and
W. A. Eaton for assistance during part of this work.
This investigation was aided by U.S. Public Health Service
grant H2520 and an American Cancer Society Institutional grant
No. IN 38D. *
?
Meyerhof, 0., Pfliiger’s Archives, 182, 233 (1920).
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7
Fleckensfcein, A., Janke, J., Davies, R. E., and Krebs, H. A., Nature,
N EWS an
Controller of Rubber Research in Malaya:
Sir Geoffrey Fletcher Clay, K.C.M.G., O.B.E,
SIR GEOFFREY FLETCHER CLAY has recently retired from the
post of controller of rubber research in Malaya, a position
which he has held since 1958, Sir Geoffrey, a graduate of the
Universities of Edinburgh and Cambridge, is well known as
an agriculturalist. After the First World War he joined the
Colonial Agricultural Service, spending many years in
Uganda, where, in 1939, he was appointed director of
agriculture. From 1942 until 1944 he was vicechairman of the
South African Production and Supply Council and from 1944
until 1946 development adviser for Northern Rhodesia and
Nyasaland. For ten years, until 1956, he was agricultural
adviser to the Secretary of State for the Colonies. Among his
many other activities, Sir Geoffrey has been a member
d VIEWS
of the United Kingdom delegation to the conference organized
by the United Nations on “Conservation of Resources” held at
Lake Success in 1949. In 1952 he led the mission sent by the
United Nations to Korea to prepare a five-year plan for the
rehabilitation of Korean agriculture, forestry and fisheries. Sir
Geoffrey has worked with the Food and Agriculture
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Organization on the problems of the reafforestation of the
Mediterranean area, and has been closely associated with the
Scientific Advisory Committee of the Empire Cotton Growing
Association.
Dr. L. C. Bateman
DB. LESLIE CLIFFORD BATEMAN has been appointed to succeed
Sir Geoffrey Clay as controller of rubber research in Malaya.
Dr. Bateman took up his position on August i and is now based
in Kuala Lumpur. As controller he is responsible for the overall
direction