Shouting strengthens voluntary force during sustained maximal effort through enhancement of motor system state via motor commands.

Previous research indicates that shouting during momentary maximal exertion effort potentiates the maximal voluntary force through the potentiation of motor cortical excitability. However, the muscular force-enhancing effects of shouting on sustained maximal force production remain unclear. We investigated the effect of shouting on the motor system state by examining motor evoked potentials in response to transcranial magnetic stimulation applied over the hand area of the contralateral primary motor cortex (M1) during sustained maximal voluntary contraction, and by assessing handgrip maximal voluntary force. We observed that shouting significantly increased handgrip maximal voluntary force and reduced the silent period. Our results indicate that shouting increased handgrip voluntary force during sustained maximal exertion effort through the reduced silent period. This is the first objective evidence that the muscular force of shouting during maximal force exertion is associated with the potentiation of motor system activity produced by the additional drive of shouting operating on the motor system (i.e., shouting-induced excitatory input to M1).

The unconscious mental inhibiting process of human maximal voluntary contraction.

Human maximal voluntary contraction (MVC) is believed to be limited by neural inhibition. Motivational goal priming alters background states of the motor system, leading to enhanced MVC. However, the mechanisms that determine the constant inhibition of force exertion in the motor system remain unclear. The primary behavioural goal of MVC is maximal voluntary force exertion. The final expected or desired state of this behavioural goal is explicitly demonstrated with words related to physical exertion, such as 'maximal', irrespective of the possibility of demand-like properties in participants' minds, such as attainability and/or desirability of the goal. For the primed maximal goal state, most trial results fail to meet expectations, demonstrating negative affect that, without awareness, contributes to the mentioned inhibitory mechanism underlying MVC. We therefore speculated that the behavioural goal of MVC contributes to neural inhibitory mechanisms underlying MVC. In our study, we used a previously developed paradigm (Takarada and Nozaki in Scientific Reports 8: 10135, 2018) in which subliminal visual priming stimuli such as the physical exertion-related words "perform" and "exert" were presented to 12 healthy participants and were followed by supraliminal words that were the word "maximal" or neutral.We found that when combined with the term 'maximal' in the consciously visible form, the effect of this subliminal motor-goal priming in inducing pupil dilation and stronger action preparation/execution was abolished without conscious awareness. This is the first objective evidence of motor inhibitory effect-predicting patterns of pupil-linked noradrenergic activity as a signature of a type of mental inhibition underlying the MVC behavioural goal.

Shouting strengthens maximal voluntary force and is associated with augmented pupillary dilation.

Previous research has demonstrated that human maximal voluntary force is generally limited by neural inhibition. Producing a shout during maximal exertion effort enhances the force levels of maximal voluntary contraction. However, the mechanisms underlying this enhancement effect on force production remain unclear. We investigated the influence of producing a shout on the pupil-linked neuromodulatory system state by examining pupil size. We also examined its effects on the motor system state by examining motor evoked potentials in response to transcranial magnetic stimulation applied over the contralateral primary motor cortex, and by evaluating handgrip maximal voluntary force. Analysis revealed that producing a shout significantly increased handgrip maximal voluntary force, followed by an increase in pupil size and a reduction of the cortical silent period. Our results indicate that producing a shout increased handgrip maximal voluntary force through the enhancement of motor cortical excitability, possibly via the enhancement of noradrenergic system activity. This study provides evidence that the muscular force-enhancing effect of shouting during maximal force exertion is related to both the motor system state and the pupil-linked neuromodulatory system state.

Motivational goal-priming with or without awareness produces faster and stronger force exertion.

Previous research has demonstrated that barely visible (subliminal) goal-priming with motivational reward can alter the state of the motor system and enhance motor output. Research shows that these affective-motivational effects result from associations between goal representations and positive affect without conscious awareness. Here, we tested whether motivational priming can increase motor output even if the priming is fully visible (supraliminal), and whether the priming effect occurs through increased cortical excitability. Groups of participants were primed with either barely visible or fully visible words related to effort and control sequences of random letters that were each followed by fully visible positively reinforcing words. The priming effect was measured behaviourally by handgrip force and reaction time to the grip cue after the priming was complete. Physiologically, the effects were measured by pupil dilation and motor-evoked potentials (MEPs) in response to transcranial magnetic stimulation during the priming task. Analysis showed that for both the supraliminal and subliminal conditions, reaction time decreased and total force, MEP magnitude, and pupil dilation increased. None of the priming-induced changes in behaviour or physiology differed significantly between the supraliminal and the subliminal groups, indicating that implicit motivation towards motor goals might not require conscious perception of the goals.

The impact of aerobic exercise training with vascular occlusion in patients with chronic heart failure.

AIMS: This study aimed to evaluate the impact of aerobic exercise training with vascular occlusion in patients with chronic heart failure. METHODS AND RESULTS: Thirty patients with post-infarction heart failure were randomized to an interventional exercise group (IG; n = 15) or a control exercise group (CG; n = 15). Exercise was performed at an intensity of 40-70% of the peak VO2 /W for 6 months. Patients in the IG remained seated on the saddle of the cycle ergometer with their feet on the pedals. Pneumatic tourniquets were applied to the proximal ends of their thighs with appropriate pressure resulting in a 40-80 mmHg increase in the systolic blood pressure that is required for vascular occlusion (208.7 ± 7.4 mmHg). We evaluated the safety and efficacy of the intervention and its effect on exercise capacity and serum BNP levels. There were no significant differences between the IG and CG in patient characteristics at study entry. Peak VO2 /W in the IG significantly increased compared with that in the CG; the change in the serum BNP levels was significantly larger in the IG than in the CG. CONCLUSIONS: These results suggest that aerobic exercise training with vascular occlusion can improve exercise capacity and serum BNP levels in patients with chronic heart failure.

Pupil dilations induced by barely conscious reward goal-priming.

The topic of unconscious influences on behavior has long been explored as a way of understanding human performance and the neurobiological correlates of intention, motivation and action. Previous research using transcranial magnetic stimulation has demonstrated that barely visible priming of an action concept, when combined with reward in the form of a consciously perceived positive stimulus, can alter the state of the motor system and enhance the maximal voluntary force level. One possible explanation is that positive stimulus-induced reward signals are processed by the dopaminergic system in the basal ganglia, motivating individuals to increase the effort they invest in particular behaviors, or to recruit the resources necessary for maintaining those behaviors. If so, given that the dopaminergic system has functionally and anatomically close connections with the noradrenergic system, we hypothesize that the state of the noradrenergic system may be enhanced by the same process. In accord with this hypothesis, we observed that barely visible goal priming with reward caused pupil dilation, suggesting that activity in the noradrenergic system increased. Importantly, this enhancement was accompanied by an unconscious increase in handgrip force. This is the first objective evidence that the pupil-linked neuromodulatory system is related to implicit learning of the link between physical exertion and reward, probably in the noradrenergic system, resulting in more forceful voluntary motor action in the absence of conscious awareness.

Maximal voluntary force strengthened by the enhancement of motor system state through barely visible priming words with reward.

The topic of unconscious influences on behaviour has long been explored as a means of understanding human performance and the neurobiological correlates of intention, motivation, and action. However, what is relatively unknown is whether subconsciously delivered priming stimuli, with or without rewards, can affect individuals' maximum level of force produced with their best effort. We demonstrated using transcranial magnetic stimulation that barely visible priming of an action concept, when combined with a reward in the form of a consciously visible positive stimulus, could alter the state of the motor system. In accordance with this neurophysiological alteration, the prime-plus-reward stimuli significantly increased the hand-grip force level of maximum voluntary contraction with little conscious awareness. This is the first objective evidence that the barely conscious presence of a behavioral goal can influence the state of the motor system and arouse latent ability for human force exertion.

Inhibition of the primary motor cortex can alter one's "sense of effort": effects of low-frequency rTMS.

Studies using force-matching tasks have suggested that when we feel a "sense of effort," cortical regions may act to increase motor commands, and thus recruit additional motor units, in order to compensate for the exerted force. We hypothesized that suppressing activity in the primary motor cortex (M1), which is the source of the motor commands, would initiate the same process, and induce the same sense of effort. In a force-matching task, grip force was applied to 'right' hand and 10 healthy participants were asked to try to exert the same amount by using 'left' hand, with no visual feedback. On some trials, low-frequency, repetitive transcranial magnetic stimulation (lf-rTMS) was used to suppress the M1 and the primary somatosensory cortex (SI) in the left hemisphere, separately. Results showed that participants tended to overestimate the level of exerted force by up to 24%. In contrast, sham stimulation of the M1 and lf-rTMS over the SI did not significantly affect participants' estimations. Further, the M1 suppression resulted in a 42% reduction in motor-evoked potentials. Thus, the M1 suppression can affect our sense of effort, suggesting that compensatory neural mechanisms that increase the MI activity may play an important role in producing senses of effort.

Hypnotic suggestion alters the state of the motor cortex.

Hypnosis often leads people to obey a suggestion of movement and to lose perceived voluntariness. This inexplicable phenomenon suggests that the state of the motor system may be altered by hypnosis; however, objective evidence for this is still lacking. Thus, we used transcranial magnetic stimulation of the primary motor cortex (M1) to investigate how hypnosis, and a concurrent suggestion that increased motivation for a force exertion task, influenced the state of the motor system. As a result, corticospinal excitability was enhanced, producing increased force exertion, only when the task-motivating suggestion was provided during hypnotic induction, showing that the hypnotic suggestion actually altered the state of M1 and the resultant behavior.

Effect of transient vascular occlusion of the upper arm on motor evoked potentials during force exertion.

We previously observed that transient vascular occlusion in volunteers increased the estimation of force exertion with no change in peripheral nerves or muscles. We hypothesized that the primary factor responsible for the overestimation of force exertion during occlusion was the centrally generated motor command, as hypothesized by McCloskey et al. (1974) and McCloskey (1978, 1981). In the present study, we tested the hypothesis that transient vascular occlusion increases the excitability of the primary motor cortex (M1) during force exertion. Healthy human volunteers lay on a bed and squeezed a dynamometer in their right hand. Repetitive gripping forces were exerted at 20%, 40%, or 60% of maximum force, with or without transient (20s) vascular occlusion of the proximal portion of the right upper arm. During the task, single-pulse transcranial magnetic stimulation was applied to the contralateral M1 to induce motor evoked potentials (MEPs) in the flexor carpi ulnaris (FCU) muscle. The MEP amplitudes were enhanced with occlusion under all conditions, with the exception of 60% contraction. In contrast, no significant difference was observed between the MEP amplitudes obtained from the occluded or non-occluded, relaxed FCU muscle. These results suggest that transient vascular occlusion increases the excitability of M1 only during force exertion.

Force overestimation during tourniquet-induced transient occlusion of the brachial artery and possible underlying neural mechanisms.

A vascular occlusion by a tourniquet inflated at the proximal end of the upper arm is suggested to affect the estimation of exertion force level. In the first part of this study, subjects were asked to estimate the isometric force exerted by the occluded hand with that of the other hand (matching experiment). We found that the perceived force with arterial occlusion was always overestimated. To examine the underlying neural mechanism for this phenomenon, in the second part, the somatosensory evoked potentials (SEPs) and nerve action potential (NAP) were recorded following electrical median nerve stimulation with or without arterial occlusion. Moreover, the maximum motor response (M response) to median nerve stimuli at the axilla was recorded from the skin surface of the thenar eminence muscle of the hand during with arterial occlusion. The N20 of SEP and NAP at Erb's point were unaffected by the arterial occlusion, and the M response was also unchanged. These results suggest that the tourniquet-induced transient occlusion of the brachial artery does not seriously affect median nerve function. Thus, it is likely that the primary responsible factor for the overestimation of perceived force exertion during arterial occlusion is the centrally generated motor command as previously hypothesized by McCloskey [McCloskey, D.I., Ebeling, P., Goodwin, G.M., 1974. Estimation of weights and tensions and apparent involvement of a "sense of effort". Exp Neurol. 42, 220-232; McCloskey, D.I., 1978. Kinesthetic sensibility. Physiol. Rev. 58, 763-820; McCloskey, D.I., 1981. Corollary discharge and motor commands and perception. In: Brookhart, J.M., Mountcastle, V.B. (Eds.), Handbook of Physiology. American Physiological Society, Bethesda, pp. 1415-1447].

Cooperative effects of exercise and occlusive stimuli on muscular function in low-intensity resistance exercise with moderate vascular occlusion.

To obtain insight into the relative contributions of exercise and occlusive stimuli to these muscular adaptations, the present study investigated the short- and long-term effects of varied combinations of low-intensity exercise and vascular occlusion. The subjects were separated into 3 groups (n = 6 for each group): low-intensity with vascular occlusion (LIO), low-intensity without vascular occlusion (LI), and vascular occlusion without exercise (VO). LIO and LI groups performed bilateral knee extension exercises in seated positions with an isotonic extension machine. In the LIO group, both sides of the thigh were pressure-occluded at the proximal end by means of a tourniquet during the entire session of exercise (approximately 10 min), whereas only the occlusion with the same pressure and duration was given in the VO group. The mean occlusion pressure was 218 +/- 8.1 mmHg (mean +/- SE). The exercise session consisted of five sets of exercise at an intensity of 10-20% 1RM and was performed twice a week for 8 wk. After the period of exercise training, isometric and isokinetic strengths at all velocities examined increased significantly in the LIO group (p < 0.05), whereas no significant change in strength was seen in the LI and VO groups. The increase in muscular strength in LIO was associated with a significant increase in the cross-sectional area of knee extensor muscles by 10.3 +/- 1.6%. The plasma growth hormone concentration measured 15 min after the session of exercise showed a marked increase only in LIO. The results showed that the low-intensity exercise and occlusive stimuli have cooperative effects in the long-term adaptation of muscle and an acute response to growth hormone.

Effects of resistance exercise combined with vascular occlusion on muscle function in athletes.

The effects of resistance exercise combined with vascular occlusion on muscle function were investigated in highly trained athletes. Elite rugby players (n = 17) took part in an 8 week study of exercise training of the knee extensor muscles, in which low-intensity [about 50% of one repetition maximum] exercise combined with an occlusion pressure of about 200 mmHg (LIO, n = 6), low-intensity exercise without the occlusion (LI, n = 6), and no exercise training (untrained control, n = 5) were included. The exercise in the LI group was of the same intensity and amount as in the LIO group. The LIO group showed a significantly larger increase in isokinetic knee extension torque than that in the other two groups (P < 0.05) at all the velocities studied. On the other hand, no significant difference was seen between LI and the control group. In the LIO group, the cross-sectional area of knee extensors increased significantly (P < 0.01), suggesting that the increase in knee extension strength was mainly caused by muscle hypertrophy. The dynamic endurance of knee extensors estimated from the decreases in mechanical work production and peak force after 50 repeated concentric contractions was also improved after LIO, whereas no significant change was observed in the LI and control groups. The results indicated that low-intensity resistance exercise causes, in almost fully trained athletes, increases in muscle size, strength and endurance, when combined with vascular occlusion.

Effects of low-intensity resistance exercise with short interset rest period on muscular function in middle-aged women.

We investigated the effect of low-intensity resistance exercise training on muscular size and strength where the interset rest period was shortened so as to reduce the metabolite clearance. Female subjects (aged 45.4 +/- 9.5 years, n = 10) performed bilateral knee extension exercises in a seated position on an isotonic leg extension machine. The exercise sessions consisted of 3 sets of exercise at a mean intensity of approximately 50% 1RM with an interset rest period of 30 seconds and was performed twice a week for a period of 12 weeks. The strength and the cross-sectional area (CSA) of the knee extensors and flexors were examined with an isokinetic dynamometer and magnetic resonance imaging (MRI), respectively. The CSAs of the knee extensors and flexors increased by 7.1 +/- 1.6% (p < 0.01, Wilcoxon signed rank test) and 2.5 +/- 1.4% (not significant), respectively. Isometric and isokinetic strengths increased significantly (p < 0.01) at all velocities examined, whereas no significant change was observed in those of knee flexors. These results indicate that a low-intensity resistance exercise with a short interset rest period is substantially effective in inducing muscular hypertrophy and concomitant increase in strength.