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[Muscle synergy in isometric flexion of the elbow].

A technique was developed for calculating the torque generated by two individual muscles (biceps brachii and brachioradialis) that contribute to the isometric flexion of the elbow. The external torque is the sum of individual torques which are unknown. Each individual torque (CB or CBR) can be related to the corresponding integrated surface EMG (QB or QBR) by means of coefficients (pB or PBR). A block of several equations C = pB QB + pBR QBR is obtained by exploring several experimental conditions. In these conditions, isometric flexion efforts of the elbow were associated to isometric efforts of supination or pronation so as to vary integrated EMG by reciprocal inhibition. By means of a least square method it was possible to know the coefficients PB and PBR. With these coefficients, it was possible to calculate the individual torques generated by the biceps brachii and brachioradialis muscles in each experimental condition.

Arm

Correction of metabolic acidosis by the kidney during isometric expansion of extracellular fluid volume.

In dogs with chronic hypochloremic metabolic alkalosis associated with ECFV contraction, plasma [HCO-3] ([HCO-3]p) normalizes during expansion of ECFV with a solution containing Cl- and HCO-3 in concentrations duplicating those in the plasma before expansion (isometric expansion). The kidney selectively rejects administered HCO-3 and retains Cl-. If this preferential Cl- less than HCO-3 reabsorptive selectivity were a characteristic renal response to ECFV expansion, isometric expansion during hyperchloremic acidosis would exacerbate the acid-base disturbance rather than correct it as it does in alkalosis. We examined the effect of isometric expansion in dogs with chronic hyperchloremic metabolic acidosis induced by HCl feeding or mineralocorticoid hormone deficiency. During expansion, as the expected decrease occurred in the fractional reabsorption of Na+, a lesser decrease occurred in fractional reabsorption of HCO-3, whereas a greater decrease occurred in fractional reabsorption of Cl-. The kidney selectively retained administered HCO3 and rejected Cl-. [HCO-3]p normalized. The shift to bicarbonate-selective from chloride-selective anion reabsorption during ECFV expansion in metabolic acidosis vs. metabolic alkalosis indicates that in response to ECFV expansion- the kidney selectively alters the ratio of bicarbonate to chloride concentration in the tubular reabsorbate in the direction that tends to normalize plasma acid-base composition, irrespective of the direction of deviation of the initial plasma bicarbonate concentration. The signal that initiates the shift in anion reabsorptive selectivity remains to be identified.

Acidosis

Effect of isometric exercise on catecholamines in the coronary circulation.

Arterial and coronary sinus blood levels of catecholamines, adenosine 3', 5'-cyclic monophosphate (c-AMP) and lactate were measured during isometric exercise in fourteen patients. In no patient did lactate production occur. Mean resting total catecholamine levels both arterial (0.53 +/- 0.07 ng/ml; 2.94 +/- 0.38 nmol/l) and coronary sinus (0.4 +/- 0.08 ng/ml; 2.22 +/- 0.44 nmol/l), did not change significantly on exercise. Coronary sinus c-AMP levels fell on exercise from 11.5 +/- 0.8 nmol/l (resting) to 9.9 +/- 0.8 nmol/l (exercise) (P less than 0.01) with an arterial-coronary sinus difference of 1.2 nmol/l (P less than 0.01) on exercise. Our findings suggest that isometric exercise does not normally result in excessive cardiac symphathetic activity.

Adult

Six weeks of isometric resistance training led to evidence of corticospinal but not reticulospinal adaptation in previously untrained adult males.

The latest hypothesis regarding the source of enhanced neural activation from resistance training is the reticulospinal rather than the corticospinal tract, based on invasive animal and emerging human data. The present study employed a six-week isometric resistance training intervention in a randomized controlled design to address this knowledge gap. Thirty-nine healthy, untrained males (age ~23 y, sustained contraction group n = 13, explosive contraction group n = 9, control group n = 17) underwent neuromuscular and electrophysiological testing and completed all study requirements. Maximal isometric torque (MVC) and rate of torque development (RTD) were measured during a familiarization session as well as before and after the six-week period. Transcranial magnetic stimulation was used to assess motor-evoked potential (MEP) area and silent period duration while subjects contracted to 10% of MVC. Loud sound (120&#xa0;dB) was used to modulate MEP area and reaction time to visual stimuli during the StartReact test. Only the intervention groups demonstrated significant improvements in MVC (27%) and RTD (60%) (both P < 0.01), along with reduced MEP area (-&#xa0;21%) and silent period duration (-&#xa0;23%) (both P < 0.01). The sustained contraction group showed reduced modulation of reaction time and increased MEP suppression due to loud sound. Short-term resistance training seemed to reduce cortical inhibition and corticospinal excitability in both training groups. The study showed conflicting changes in measures purported to evaluate reticulospinal functioning. It is recommended to examine different forms of resistance training and longer training exposure in future.

Humans

Effect of isometric exercises on body potassium and dietary protein requirements of young men.

A diet supplying 0.5 or 1 g egg and milk protein/kg per day and adequate energy for maintenance of body weight was given to seven healthy men aged 18 to 21. After 2 weeks for adaptation to the diet they participated in 75 min of daily isometric exercises for 4 to 6 weeks. Two men on each protein intake continued to exercise for 4 to 5 additional weeks after undergoing changes in their daily protein intake from 0.5 to 1 g/kg or vice versa. A progressive decrease in total body potassium became significant by the fourth week of exercises in all men who began on the 0.5 g protein diet. Two men on that diet who remained as nonexercising controls lost no body K. These findings indicated that 0.5 g of egg and milk protein/kg per day was insufficient for men performing isometric exercises without prior training, whereas 1 g/kg per day appeared to be adequate. Two other nonexercising men on the 0.5 g diet who lost weight showed a marked negative nitrogen balance and loss of total body potassium which improved after their dietary energy intake increased. The changes in nitrogen balance and body weight observed in this investigation support the concept that protein requirements vary with energy intakes. Conversely, they suggest that energy requirements are influenced by the level of protein intake.

Adult

Effect of adrenergic receptor blockade on the responses to isometric handgrip: studies in normal and hypertensive subjects.

Heart rate and blood pressure (BP) responses to a standardized 3 min isometric handgrip (IHG) test were measured in 32 normotensive men and compared with those found in 35 age-matched, drug-free men with established essential hypertension (BP range: 140--170/90--110). IHG testing in the normal subjects induced significant increases in heart rate (mean +/- SE: 7.6 +/- 3.3 beats/min), systolic blood pressure (19.4 +/- 5.3 mm Hg), and diastolic blood pressure (15.6 +/- 4.6 mm Hg). Although beginning from higher resting levels the hypertensive patients had similar degrees of increase in all three parameters. After chronic treatment with propranolol, the heart rate increase with IHG was suppressed in both study groups, but blood pressure responses differed, with a diminished pressor response to IHG seen in normal subjects and augmented pressor effects in the hypertensive group. Intravenous administration of phentolamine and propranolol completely abolished the pressor effects of IHG. These observations suggest that the autonomic control mechanisms mediating the responses to isometric exercise function similarly in drug-free normal and hypertensive patients and that the responses to IHG, mediated largely by endogenous catecholamine release, can be prevented by peripheral sympathetic receptor blockade.

Adrenergic alpha-Antagonists

Dose-response effects of isometric wall squats on postexercise hypotension in older women with hypertension.

OBJECTIVES: The isometric wall squat (IWS) is a promising nonpharmacological intervention for blood pressure (BP) management, but its optimal dosage in older adults remains unclear. This study compared postexercise hypotension (PEH) after two IWS doses in medicated older women with systemic arterial hypertension (SAH). METHODS: Thirty older hypertensive women (68.8&#x200a;&#xb1;&#x200a;10.0&#x200a;years) were randomized into three groups: control (CG), 8-min IWS (G8-IWS; 4&#x200a;&#xd7;&#x200a;2-min sets), and 12-min IWS (G12-IWS; 6&#x200a;&#xd7;&#x200a;2-min sets). BP and heart rate (HR) were assessed at rest and at 0, 5, 10, 15, 30, 45, and 60&#x200a;min postexercise and analyzed using linear mixed-effects models with Bonferroni correction ( P &#x200a;<&#x200a;0.05). RESULTS: G12-IWS significantly reduced systolic BP and mean arterial pressure (MAP) at 45-60&#x200a;min and 30-60&#x200a;min postexercise ( P &#x200a;<&#x200a;0.05), while G8-IWS showed no BP changes. G12-IWS decreased HR (30-60&#x200a;min) and rate-pressure product (RPP; 15-60&#x200a;min); G8-IWS reduced RPP from 30 to 60&#x200a;min only. Diastolic BP and pulse pressure remained unchanged in both groups. CONCLUSION: A single G12-IWS session induced clinically meaningful PEH (~8-10&#x200a;mmHg), exceeding reported with isometric handgrip exercise (~4-6&#x200a;mmHg), with greater RPP suppression than G8-IWS. This equipment-free protocol represents a relevant nonpharmacological strategy for BP management in older women with SAH.

Humans

Influence of temperature on isometric contraction and passive muscular tension in paramyotonia congenita (Eulenburg).

Four patients without symptoms of episodic hyperkalemic weakness from two families with paramyotonia congenita (Eulenburg) are described. 1. Maximum voluntary muscle contraction of the upper and lower arm was studied under isometric conditions at different temperatures. If the temperature was lowered stepwise, distinct paresis occured at 32--31 degrees C which increased with the amount of muscular effort. The upper arm muscles, however, developed weakness gradually after cooling. 2. During cooling of the resting muscle, the EMG showed dense spontaneous activity of the fibrillary type, which decreased again at about 30 degrees C. It can be assumed that in paramyotonia congenita cooling produces muscle cell membrane depolarization which at a critical level causes the firing of action potentials and finally muscular paresis. 3. Increasing muscular stiffness can be interpreted as abnormally slow muscular relaxation after isometric contraction. In the forearm muscles the time to 3/4 relaxation after cooling was about six times normal, in the upper arm muscles only two times normal. As an additional parameter the mechanical resistance to passive stretching of a muscle has been studied. This passive muscular tension increased simultaneously with the onset of weakness. 4. The close relation between weakness and stiffness suggest that both symptoms are caused by the same basic defect which is probably located in the sarcolemma. It is suggested that a defect of the sodium channel causes a cooling-dependent increase in sodium conductance. Raised intracellular sodium causes in the first place membrane depolarization, and in the second place depression of calcium reuptake through competition by sodium for calcium binding sites. This would explain muscle stiffness and delayed relaxation as well.

Arm

The relationship between the rate of rise of isometric tension and motor unit recruitment in a human forearm muscle.

The recruitment properties of single motor units from the human extensor indicis muscle were investigated during voluntary isometric contractions of different rate of rise but equal amplitude. Both the electrical and the contractile events associated with recruitment were analyzed. The threshold force of recruitment (measured as the total muscle force at firing onset) decreased with increasing rate of rise of isometric tension. This was consistently found for all units. Differences between low and high threshold units indicating a preferential tonic or phasic mode of activation were not observed. The contractile events associated with recruitment were analyzed analoguously to the electrical events. For this purpose, muscle force was measured at the time of the first twitch as it was measured at the time of the first spike. This separate measurement of the electrical and mechanical recruitment of a unit is necessary, because during a change of muscle force, force is different at firing onset and during the subsequent twitch contraction. Muscle tension at the time of the peak of the first twitch contraction was calculated from measurements of the twitch concentration time of the single motor units. In contrast to firing onset, the peak of the first twitch of a motor unit occurs at approximately the same muscle tension no matter how fast the contraction is performed. This is the consequence of the result that the average decrease of the threshold force of recruitment at successively faster contraction has the same value as the corresponding increase of total muscle force during the mean contraction time of the motor units. On the basis of this precise matching between these two changes, the mechanical recruitment of motor units occurs at approximately the same force level irrespective of the rate of rise of tension.

Electric Stimulation

Analytical approach to mathematical modelling of the isometric contraction of the hypertrophied myocardium.

It is attempted on the basis of the Hill two-element model to simulate mathematically the rising phase of the isometric mechanogram in the pressure-hypertrophied rat myocardium (Goldblatt II). The evaluation is based on the experimentally determined force-velocity relation as well as on the characteristic of the series-elastic stiffness. The changes in the isometric mechanogram of the hypertrophied heart muscle observed in the experiment can be mathematically predicted solely by means of the alterations in the force-velocity relationships induced by the hypertrophy, whereby the characteristic of the series-elastic stiffness is considered to be unchanged.

Cardiomegaly

Influence of low extracellular pH upon the Ca inward current and isometric contractile force in mammalian ventricular myocardium.

In isolated papillary muscles of cats the changes in Ca inward current and isometric contractile force following a decrease of extracellular pH from 7.4 to 5.5 were studied. The Ca current was analyzed (a) by measuring the upstroke velocity of Ca-mediated action potentials and (b) in voltage clamp experiments using the double sucrose gap technique. 1. At a pH of 5.5 the upstroke velocity of the Ca-mediated action potential decreased to 65% of the control, while overshoot and action potential duration remained almost unchanged. Furthermore, the relative refractory period was prolonged and in some cases, a "Wenckebach-like" phenomenon occurred. In voltage clamp experiments, the slow inward current was found to be diminished to 50-60% of the initial control value and over a broad voltage range the current voltage relationship curve was shifted to weaker currents. Acidosis did not influence the steady state inactivation but altered the kinetics of inactivation of the slow inward current and induced an increase of tauinactivation and taurecovery. This indicates that acidosis exerts a complex effect on the slow membrane channel. 2. The normal response of the Ca current towards variations of the extracellular Ca concentration (0.5-4 mM) or towards the addition of the beta-stimulating compound isoproterenol (2 mg/l) was not altered by the lowered extracellular pH. 3. In the acid medium, isometric contractile force declined to 40% of the control value within 25 min and, thus, reacted stronger than the Ca current. This indicates that those forms of acidosis used in the present experiments caused their negative inotropic effect not exclusively via a depression of the Ca current. Rather an additional intracellular effect has to be assumed which finally leads to a reduced activity of the contractile system. 4. At pH 5.5 excess Ca (4 mM) induced the same quantitative response of the contractile system as obtained at normal pH. In contrast, the positive-inotropic effect of 2 mg/l isoproterenol was more pronounced, whilst the sensitivity of the Ca inward current towards this beta-stimulating compound remained unchanged.

Action Potentials

The time-course of energy balance in an isometric tetanus.

Unpoisoned sartorius muscles of Rana temporaria were stimulated tetanically in isometric contractions lasting up to 20 s at 0 degrees C. The observed enthalpy (heat + work) production and the chemical changes in these contractions were measured, and a comparison was made between the observed enthalpy and the enthalpy that could be explained by the chemical changes. Like earlier workers, we found that the only net known reaction of energetic significance that occurred was dephosphorylation of n-phosphoryl creatine (PC), and we found a significant evolution of unexplained enthalpy (UE), a portion of the observed enthalpy which could not be explained by the extent of PC dephosphorylation. We measured the total quantity and the rate of production of the UE, and we found that its rate of evolution, which was most rapid during the first 750 ms of contraction, fell progressively to zero by the 8th s of contraction: i.e., after 8 s of contraction, all the observed enthalpy is adequately explained by PC dephosphorylation. The time-course of evolution of the UE was slower than that of the labile enthalpy (a component of the enthalpy evolved in isometric contraction whose rate of production declines exponentially at approximately 1 s-1). We conclude that, although the magnitudes of these enthalpy quantities may be similar, they are not derived from the same chemical reaction in muscle.

Animals

The responses of human muscle spindle endings to vibration during isometric contraction.

1. An human subjects, vibration of amplitude 1-5 mm and frequency 20-220 Hz was applied to the tendons of muscles in the leg to examine the effects on the discharge of primary and secondary endings during manoeuvres designed to alter the level of fusimotor drive. 2. In four experiments, the peroneal nerve was completely blocked with lidocaine proximal to the recording site in order to de-efferent spindle endings temporarily. The responses to muscle stretch and vibration, as seen in multi-unit recordings and in single unit recordings, were similar during the block as in the relaxed state prior to the block. Thus, these experiments provided no evidence for a functionally effective resting fusimotor drive. 3. The responses to vibration of nine primary endings and four secondary endings were examined during isometric voluntary contractions of the receptor-bearing muscles. Providing that the endings were responding submaximally in the relaxed state, voluntary contraction enhanced the response to vibration, suggesting co-activation of the fusimotor system sufficient to compensate for mechanical unloading. Unloading effects were observed during contractions of neighbouring synergistic muscles, indicating a close spatial relationship between the co-activated skeletomotor and fusimotor outflows. 4. Recordings were obtained from ten primary endings and seven secondary endings during isometric reflex contractions resulting from the vibratory stimulus (TVR contractions). For twelve endings, the appearance of the tonic vibration reflex in the receptor-bearing muscle resulted in a significant decrease in the response to vibration, suggesting that the endings were unloaded by the extrafusal contraction. On voluntary suppression of the reflex contraction spindle responses reverted to their previous levels. 5. These results suggest that the tonic vibration reflex, like the tendon jerk reflex, operates predominantly or exclusively on alpha motoneurones and that it does not utilize the same cortically originating efferent pathways as are used in the performance of voluntary contractions.

Action Potentials

Recruitment order of human spindle endings in isometric voluntary contractions.

1. The responses of twenty-two spindle endings in the anterior tibial and toe extensor muscles of human subjects were studied during isometric voluntary contractions of the receptor-bearing muscle with the ankle joint fixed in 25 degrees plantar flexion.2. The discharge of eighteen endings accelerated in voluntary contractions when the contraction strength exceeded a threshold level which differed for different endings but was reproducible for the same ending.3. With contractions of slow onset the latency to spindle acceleration varied with the speed of onset of the contraction. Endings with a background discharge were often unloaded by contractions until the contraction strength exceeded the threshold for activation of the ending.4. No correlation was found between the sensitivity of a spindle to external length changes and its ease of activation in a voluntary contraction. For two spindle endings with a background discharge there was no change in either discharge frequency or the regularity of spindle discharge during contractions which were below the threshold for activation of the endings. It is concluded that the threshold for activation of a spindle ending in an isometric voluntary contraction is determined by its fusimotor innervation, and that fusimotor neurones probably have a recruitment order, much as do skeletomotor neurones.5. Once activated, the discharge of spindle endings fluctuated with changes in skeletomotor activity but the relationship for some endings contained non-linearities. Such non-linearities were not as apparent in multi-unit recordings from a number of spindle endings in the contracting muscle. It is concluded that the fusimotor drive to a muscle is proportional to the skeletomotor drive to the muscle, and that skeletomotor and fusimotor neurones are subjected to similar if not identical descending command signals. From the work of Evarts (1968), it seems likely that these command signals are related more to desired muscle force than to desired muscle length.

Action Potentials

Force-velocity relation in normal and nitrate-treated frog single muscle fibres during rise of tension in an isometric tetanus.

1. The force-velocity (P-V) relation for normal or NO-3 treated single fibres isolated from the semitendinosus muscle of the frog was determined at given times during the rise of tension and the plateau of isometric tetani. Experiments were made at about 2.25 micron sarcomere length and at constant temperatures, from 3 to 4.5 degrees C and from 19 to 21 degrees C. The controlled-velocity release method was used. 2. During the rise of tension, at any initial tension higher than about 0.2 P0, the lowest release velocity required to drop the tension to zero was the same as at the tetanic plateau, independent of the temperature and the presence of NO-3 ions in the bathing solution. 3. The degree of activation (measured by the steady force exerted at a given velocity of shortening lower than V0) increased with time, but attained its steady-state level before isometric tension. 4. At about 20 degrees C, frog muscle fibres at about 2.2 micron sarcomere length were only partially activated after a single stimulus. 5. NO-3 ions did not affect the steady-state P-V relation. At about 20 degrees C, NO-3 ions increased the rate of development of activation. Potentiation of the twitch contraction was due at least in part to this mechanism. 6. The 'relative' P-V relation appears to be independent of both the time after start of stimulation and the presence of NO-3 ions in the bathing solution. 7. The results are discussed in terms of the sliding filament model of Huxley (1957), assuming that either the number of actin sites available for cross-bridge formation, or the value of the rate constant for making of cross-bridges, is time dependent.

Animals

The velocity of unloaded shortening and its relation to sarcomere length and isometric force in vertebrate muscle fibres.

1. The velocity of shortening at zero load was studied during fused tetanic contractions and single twitches in isolated skeletal muscle fibres of Rana temporaria. 2. The technique used for determination of the speed of unloaded shortening consisted of a series of quick releases of different amplitudes applied at a given instant during activity. The time, delta t, needed for the fibre to take up the slack was plotted against the amplitude of release, delta L. The slope of the straight line relating delta t-delta L provided a measure of the velocity of shortening at zero load, V0. 3. V0 was compared with force-velocity data obtained at finite loads (load-clamp recordings). The predicted velocity of shortening at zero load, derived by hyperbolic extrapolation from velocities at low and intermediate loads, was not significantly different from V0. 4. The temperature dependence of isometric force and of shortening velocity was investigated between 2 and 12 degrees C in the same fibres. Q10 was 2.67 +/- 0.07 (S.E. of mean, n = 6) for V0 and 1.24 +/- 0.01 for tetanic force. 5. The velocity of unloaded shortening was determined at different sarcomere lengths in the range 1.4--3.1 microns. V0 was constant between 1.65 microns and approximately 2.7 microns. It decreased below 1.65 microns and increased above 2.7 microns. 6. The decrease in velocity at short sarcomere lengths probably reflects an increase of the passive resistance to shortening. The increase in velocity at long sarcomere lengths can be accounted for by the passive compressive force that is produced by the parallel elastic elements of the prestretched fibre. 7. V0 was determined at the peak of the twitch and during the plateau of the fused tetanus in the same fibre. Whereas the peak twitch force varied between 38 and 85% of the tetanic tension in the different fibres (mean: 71 +/- 5%, n = 8), V0 during the twitch was 99 +/- 2% of the value recorded during the tetanus. Depression of the isometric twitch amplitude to 10% of the control value by dantrolene did not cause any significant reduction of V0.

Animals

Tremor in the tension developed isometrically by soleus during the tonic vibration reflex in the decerebrate cat.

1. Irregularities in the development of tension during the tonic vibration reflex of the soleus muscle of the decerebrate cat have been analysed into their frequency components. The reflex was recorded isometrically and elicited by longitudinal vibration, normally at 150 Hz. The amplitude of vibration was set so as to elicit a maximal reflex response, suggesting 1:1 driving of the majority of the Ia afferents at the frequency of vibration. 2. The resulting power spectrum regularly showed a well marked tremor peak separated by a trough from any slow irregularities. The predominant frequency of this tremor varied from 4 to 11 Hz in different preparations, with a mean of 7.4 Hz; on average, frequencies within 1.7 Hz on either side contained over half the power of the predominant frequency. Altering the frequency of vibration did not alter the distribution of tremor frequencies. 3. The root mean square value of the tension irregularities, over the range 4-14 Hz, varied from 12 to 110 mN in different preparations (median value, 23 mN); this was superimposed on mean active reflex tensions varying from 2 to 10 N. 4. The 'tremor' due to a single motor unit was estimated from spectral analysis of tetanic contractions of the whole muscle and decreased with increasing frequency of activation. Comparison of the single unit values with the tremor seen during vibration in the same preparations showed that equivalent amounts of tremor to the latter could typically have been produced by the continued synchronous contraction of about five 'average' motor units firing at the predominant tremor frequency. 5. When a tonic stretch reflex was present its tremor frequencies did not differ consistently from those of the tonic vibration reflex. On average, the tremor was smaller for the stretch reflex than for the tonic vibration reflex; the difference was usually slight and might have been related to the stretch refex tension being smaller. 6. Evidence was obtained that the tremor was not due to any insecurity of 1:1 driving of the Ia afferents by the vibration. First, the tremor did not increase when the amplitude of vibration was decreased sufficiently to ensure that the degree of 1:1 driving must have been reduced. Secondly, the introduction of a comparable 'artificial tremor' by sinusoidally oscillating the muscle at low frequency did not produce the e.m.g. response that would have been expected if the applied 'tremor' had been modulating the firing of the Ia or any other group of afferents. 7. It is concluded that the observed tremor cannot be attributed to 'oscillation in the stretch reflex arc', though without prejudice to the role of this mechanism under other conditions and especially when the recording is not isometric. However, the genesis of the tremor has not been established and much of it might result simply from the chance synchronization of motor units that are firing below their tetanic fusion frequency.

Action Potentials

Constituents of human muscle in isometric fatigue.

Three subjects performed five successive isometric contractions to fatigue; the tension in any one experiment was constant at tensions varying from 20 to 80% of the maximal voluntary contraction (MVC). The interval between contractions was held constant at 11 min. Muscle biopsy specimens were obtained at the start of the experiment, after the first, fourth, and fifth, and before the second and fifth of the successive contractions. The concentrations of ATP, CP, glycogen, and lactate were measured in each sample of muscle. Changes in ATP and glycogen were insufficient to be held accountable for the development of isometric fatigue. Changes in CP and lactate were large after fatigue at intermediate tensions, but those of CP were considered unlikely to be responsible for the fatigue. At tensions of 30-50% MVC the increase in lactate could be responsible for fatigue either directly or by indirect changes in pH; at higher and lower tensions the possibility that lactate is directly implicated in the development of fatigue seems remote.

Adenosine Triphosphate