Heterogeneity of psychophysiological stress responses in fibromyalgia syndrome patients

Dysregulated psychophysiological responses have been observed in patients with fibromyalgia syndrome (FMS), although the results are inconsistent. Surface electromyographic (EMG), systolic and diastolic blood pressure, heart rate (HR), and skin conductance levels (SCLs) were continuously recorded at baseline, and during a series of stress and relaxation tasks in 90 FMS patients and 30 age and sex matched healthy controls (HCs). The patient sample demonstrated lower baseline EMG levels compared to the HCs on all tasks. In contrast, the patients displayed elevated HR and SCL (sympathetic vasomotor and sudomotor indices, respectively) during both stress tasks. A cluster analysis identified four psychophysiological response patterns: 63.3% of HCs showed increased muscle tension and stable cardiovascular responses; 34.8% of FMS patients showed a pattern of increased sympathetic vasomotor reactivity with stable sudomotor and reduced muscular response; 12.2% of FMS patients showed a pattern of increased sympathetic sudomotor reactivity connected with increased sympathetic vasomotor response and reduced muscular response; and, in contrast, 46.7% of FMS patients showed a pattern of parasympathetic vasomotor reactivity and reduced sudomotor as well as muscular response. The identification of low baseline muscle tension in FMS is discrepant with other chronic pain syndromes and suggests that unique psychophysiological features may be associated with FMS. The different psychophysiological response patterns within the patient sample support the heterogeneity of FMS.     

Heterogeneity of psychophysiological stress responses in fibromyalgia syndrome patients

Dysregulated psychophysiological responses have been observed in patients with fibromyalgia syndrome (FMS), although the results are inconsistent. Surface electromyographic (EMG), systolic and diastolic blood pressure, heart rate (HR), and skin conductance levels (SCLs) were continuously recorded at baseline, and during a series of stress and relaxation tasks in 90 FMS patients and 30 age and sex matched healthy controls (HCs). The patient sample demonstrated lower baseline EMG levels compared to the HCs on all tasks. In contrast, the patients displayed elevated HR and SCL (sympathetic vasomotor and sudomotor indices, respectively) during both stress tasks. A cluster analysis identified four psychophysiological response patterns: 63.3% of HCs showed increased muscle tension and stable cardiovascular responses; 34.8% of FMS patients showed a pattern of increased sympathetic vasomotor reactivity with stable sudomotor and reduced muscular response; 12.2% of FMS patients showed a pattern of increased sympathetic sudomotor reactivity connected with increased sympathetic vasomotor response and reduced muscular response; and, in contrast, 46.7% of FMS patients showed a pattern of parasympathetic vasomotor reactivity and reduced sudomotor as well as muscular response. The identification of low baseline muscle tension in FMS is discrepant with other chronic pain syndromes and suggests that unique psychophysiological features may be associated with FMS. The different psychophysiological response patterns within the patient sample support the heterogeneity of FMS.     

THE RELEVANT USE OF THE TRADITIONAL TUNISIAN GAME ‘‘RAQASSA’’ FOR CARDIOVASCULAR STIMULATION IN SCHOOLCHILDREN

The aim of this study was to investigate the heart rate (HR) responses, the rate of perceived exertion (RPE), and the feeling during physical education schooling while performing traditional games activities compared to intermittent exercise. Nineteen pre-pubertal children randomly performed on different days two types of lessons (intermittent running mode vs. traditional Tunisian “Raqassa” game) lasting 12-min each. HR was continuously recorded during both lessons, while ratings of perceived exertion and Feeling values were recorded after the sessions. The mean HR value during the traditional game was significantly higher than during intermittent exercise (p<0.05). Conversely, the perceived exertion score was significantly higher after intermittent exercise than the traditional exercise game (p<0.05), showing that the higher cardiovascular strain of the game was perceived as “lighter” than the run. Simultaneously, the children''s Feeling was significantly higher after the traditional game than intermittent exercise (p<0.001), showing a higher satisfaction from playing with respect to running. Exercise based on the “Raqassa” traditional game could be used in pre-pubertal children as an alternative or as an additional method for suitable cardiovascular stimulation during physical education lessons with lower perceived exertion and better feeling compared to intermittent running.     

Physiological and psychological indices of fatigue during static contractions

The development of fatigue and the relationship between psychological and physiological indices of fatigue were studied in a group of 18 male subjects during static contractions. Exercise was performed as a static elbow flexion at 25% MVC. Heart rate (HR), intraarterial blood pressure (BP) and surface EMG [mean amplitude (MA) and central frequency (CF)] were studied during contractions sustained until exhaustion. The amount of effort expended (relative to total exhaustion) and the rating of perceived pain were recorded following contractions interrupted after 20, 30 . . . 80, and 100% of endurance time. HR, BP and EMG amplitude responses were similar to those previously recorded. The decline in CF occurred in two phases, possibly related to a change in motor unit recruitment after the initial 70% of endurance time. The subjects overestimated the amount of effort expended and thus underestimated their endurance capacity. The best correlation between perceived effort and physiological responses was obtained using blood pressure data, whereas changes in EMG data did not parallel the psychological responses. It is concluded that the perception of effort during a static contraction is produced through a complex process, in which several influences of peripheral and central origin are integrated.     

Facial EMG Responses to Emotional Expressions Are Related to Emotion Perception Ability

Although most people can identify facial expressions of emotions well, they still differ in this ability. According to embodied simulation theories understanding emotions of others is fostered by involuntarily mimicking the perceived expressions, causing a “reactivation” of the corresponding mental state. Some studies suggest automatic facial mimicry during expression viewing; however, findings on the relationship between mimicry and emotion perception abilities are equivocal. The present study investigated individual differences in emotion perception and its relationship to facial muscle responses - recorded with electromyogram (EMG) - in response to emotional facial expressions. N° = °269 participants completed multiple tasks measuring face and emotion perception. EMG recordings were taken from a subsample (N° = °110) in an independent emotion classification task of short videos displaying six emotions. Confirmatory factor analyses of the m. corrugator supercilii in response to angry, happy, sad, and neutral expressions showed that individual differences in corrugator activity can be separated into a general response to all faces and an emotion-related response. Structural equation modeling revealed a substantial relationship between the emotion-related response and emotion perception ability, providing evidence for the role of facial muscle activation in emotion perception from an individual differences perspective.     

Psychophysiological Response Patterns to Affective Film Stimuli

Psychophysiological research on emotion utilizes various physiological response measures to index activation of the defense system. Here we tested 1) whether acoustic startle reflex (ASR), skin conductance response (SCR) and heart rate (HR) elicited by highly arousing stimuli specifically reflect a defensive state and 2) the relation between resting heart rate variability (HRV) and affective responding. In a within-subject design, participants viewed film clips with a positive, negative and neutral content. In contrast to SCR and HR, we show that ASR differentiated between negative, neutral and positive states and can therefore be considered as a reliable index of activation of the defense system. Furthermore, resting HRV was associated with affect-modulated characteristics of ASR, but not with SCR or HR. Interestingly, individuals with low-HRV showed less differentiation in ASR between affective states. We discuss the important value of ASR in psychophysiological research on emotion and speculate on HRV as a potential biological marker for demarcating adaptive from maladaptive responding.     

Can Cognitive Activities during Breaks in Repetitive Manual Work Accelerate Recovery from Fatigue? A Controlled Experiment

Neurophysiologic theory and some empirical evidence suggest that fatigue caused by physical work may be more effectively recovered during “diverting” periods of cognitive activity than during passive rest; a phenomenon of great interest in working life. We investigated the extent to which development and recovery of fatigue during repeated bouts of an occupationally relevant reaching task was influenced by the difficulty of a cognitive activity between these bouts. Eighteen male volunteers performed three experimental sessions, consisting of six 7-min bouts of reaching alternating with 3 minutes of a memory test differing in difficulty between sessions. Throughout each session, recordings were made of upper trapezius muscle activity using electromyography (EMG), heart rate and heart rate variability (HRV) using electrocardiography, arterial blood pressure, and perceived fatigue (Borg CR10 scale and SOFI). A test battery before, immediately after and 1 hour after the work period included measurements of maximal shoulder elevation strength (MVC), pressure pain threshold (PPT) over the trapezius muscles, and a submaximal isometric contraction. As expected, perceived fatigue and EMG amplitude increased during the physical work bouts. Recovery did occur between the bouts, but fatigue accumulated throughout the work period. Neither EMG changes nor recovery of perceived fatigue during breaks were influenced by cognitive task difficulty, while heart rate and HRV recovered the most during breaks with the most difficult task. Recovery of perceived fatigue after the 1 hour work period was also most pronounced for the most difficult cognitive condition, while MVC and PPT showed ambiguous patterns, and EMG recovered similarly after all three cognitive protocols. Thus, we could confirm that cognitive tasks between bouts of fatiguing physical work can, indeed, accelerate recovery of some factors associated with fatigue, even if benefits may be moderate and some responses may be equivocal. Our results encourage further research into combinations of physical and mental tasks in an occupational context.     

The Relation between Thematic Role Computing and Semantic Relatedness Processing during On-Line Sentence Comprehension

Sentence comprehension involves timely computing different types of relations between its verbs and noun arguments, such as morphosyntactic, semantic, and thematic relations. Here, we used EEG technique to investigate the potential differences in thematic role computing and lexical-semantic relatedness processing during on-line sentence comprehension, and the interaction between these two types of processes. Mandarin Chinese sentences were used as materials. The basic structure of those sentences is “Noun+Verb+‘le’+a two-character word”, with the Noun being the initial argument. The verb disambiguates the initial argument as an agent or a patient. Meanwhile, the initial argument and the verb are highly or lowly semantically related. The ERPs at the verbs revealed that: relative to the agent condition, the patient condition evoked a larger N400 only when the argument and verb were lowly semantically related; however, relative to the high-relatedness condition, the low-relatedness condition elicited a larger N400 regardless of the thematic relation; although both thematic role variation and semantic relatedness variation elicited N400 effects, the N400 effect elicited by the former was broadly distributed and reached maximum over the frontal electrodes, and the N400 effect elicited by the latter had a posterior distribution. In addition, the brain oscillations results showed that, although thematic role variation (patient vs. agent) induced power decreases around the beta frequency band (15–30 Hz), semantic relatedness variation (low-relatedness vs. high-relatedness) induced power increases in the theta frequency band (4–7 Hz). These results suggested that, in the sentence context, thematic role computing is modulated by the semantic relatedness between the verb and its argument; semantic relatedness processing, however, is in some degree independent from the thematic relations. Moreover, our results indicated that, during on-line sentence comprehension, thematic role computing and semantic relatedness processing are mediated by distinct neural systems.     

Heart rate responses to Taekwondo training in experienced practitioners

The purpose of this study was to evaluate the heart rate (HR) responses of specific Taekwondo training activities, practiced by experienced practitioners in a natural training environment. Eight male experienced Taekwondo practitioners, with 3- 13 years (5.4 +/- 3.2 years) experience took part in a 5-day Taekwondo training camp. Continuous HR measures were recorded at 5-second intervals during 6 training sessions; each session was observed and notated, and a diary of training activities was recorded. The HR responses were assimilated into 8 fundamental training activities for analysis: elastics, technical combinations, step sparring, pad work, forms, basic techniques and forms, sparring drills, and free sparring. Taekwondo training elicited HR into 64.7-81.4% of HR maximum (%HRmax). Moderate relative exercise intensities (64.7-69.4%HRmax) were elicited by elastics, technical combinations, and step sparring. The remaining 5 training activities elicited hard relative exercise intensities (74.7-81.4%HRmax). One-way repeated-measures analysis of variance with post hoc analysis revealed that elastics, technical combinations, and step sparring elicited significantly lower relative intensities than the remaining training activities (p < 0.05). Furthermore, forms, basic techniques and forms, sparring drills, and free sparring elicited significantly higher relative intensities than the remaining training activities (p < 0.05). In conclusion, all Taekwondo training activities in this study seemed suitable for cardiovascular conditioning, although different training activities stressed the cardiovascular system to different degrees. Practically, this suggests coaches need to structure Taekwondo training sessions based not only on the technical and tactical needs of practitioners but also in a manner that enables sufficient cardiovascular conditioning for competition.     

Psychophysiological Characteristics of Nuclear Power Station Operator Activity as a Factor in Accident Rate

Methods of psychological, physiological, and psychophysiological testing were used to assess the intensity of the activity of operators of the power unit control panel of the Khmel'nitskaya nuclear power station. The operators during whose duty serious control errors and preaccident situations were noted were examined. The psychological and psychophysiological characteristics of these operators were shown to differ in that their reactive anxiety was increased and their well-being, activity, and mood scores, as well as the functional reserves of the cardiovascular system and mental and physical working capacity, were decreased. These parameters were considered as criteria in the development of operating measures. It was proposed that not only technological factors but also operators' psychophysiological characteristics in the dynamics of their manifestation be taken into account for elaborating measures of optimal staff selection and placement.     

Regulation of mental states and biofeedback techniques: Effects on breathing pattern

The purpose of the present study was to examine whether breathing pattern may be used as a reliable index for the effectiveness of techniques applied for the regulation of mental states. Heart rate (HR), breathing pattern, galvanic skin response (GSR), and electromyogram (EMG) of the frontalis muscle were measured in 39 male and female subjects aged 18–25 years during 10-minute treatment with relaxation technique (autogenic training and/or music) followed by 10 minutes of imagery training. In the first 7 sessions biofeedback (BFB) was not included, while during the last 6 sessions BFB was introduced and utilized by the subjects. Relaxation (music or autogenic training) led to a decrease in breathing frequency, attributed to lengthening of expiration time, as well as reduced HR, GSR, and frontalis EMG response. In most instances imagery training was related to an increase in these indices. Specifically, significant tachypnea was observed during imagery of sprint running. In most cases BFB substantially augmented the physiological responses. In conclusion, our data suggest that, compared with HR, GSR, and EMG responses, the breathing pattern is at least as sensitive to the mental techniques employed, and may be useful as a psychophysiological index for diagnosis and testing, especially in sport practice.     

Relativity Theory and Time Perception: Single or Multiple Clocks?

Background

Current theories of interval timing assume that humans and other animals time as if using a single, absolute stopwatch that can be stopped or reset on command. Here we evaluate the alternative view that psychological time is represented by multiple clocks, and that these clocks create separate temporal contexts by which duration is judged in a relative manner. Two predictions of the multiple-clock hypothesis were tested. First, that the multiple clocks can be manipulated (stopped and/or reset) independently. Second, that an event of a given physical duration would be perceived as having different durations in different temporal contexts, i.e., would be judged differently by each clock.

Methodology/Principal Findings

Rats were trained to time three durations (e.g., 10, 30, and 90 s). When timing was interrupted by an unexpected gap in the signal, rats reset the clock used to time the “short” duration, stopped the “medium” duration clock, and continued to run the “long” duration clock. When the duration of the gap was manipulated, the rats reset these clocks in a hierarchical order, first the “short”, then the “medium”, and finally the “long” clock. Quantitative modeling assuming re-allocation of cognitive resources in proportion to the relative duration of the gap to the multiple, simultaneously timed event durations was used to account for the results.

Conclusions/Significance

These results indicate that the three event durations were effectively timed by separate clocks operated independently, and that the same gap duration was judged relative to these three temporal contexts. Results suggest that the brain processes the duration of an event in a manner similar to Einstein''s special relativity theory: A given time interval is registered differently by independent clocks dependent upon the context.     

Distinguishing Neurocognitive Processes Reflected by P600 Effects: Evidence from ERPs and Neural Oscillations

Research on language comprehension using event-related potentials (ERPs) reported distinct ERP components reliably related to the processing of semantic (N400) and syntactic information (P600). Recent ERP studies have challenged this well-defined distinction by showing P600 effects for semantic and pragmatic anomalies. So far, it is still unresolved whether the P600 reflects specific or rather common processes. The present study addresses this question by investigating ERPs in response to a syntactic and pragmatic (irony) manipulation, as well as a combined syntactic and pragmatic manipulation. For the syntactic condition, a morphosyntactic violation was applied, whereas for the pragmatic condition, such as “That is rich”, either an ironic or literal interpretation was achieved, depending on the prior context. The ERPs at the critical word showed a LAN-P600 pattern for syntactically incorrect sentences relative to correct ones. For ironic compared to literal sentences, ERPs showed a P200 effect followed by a P600 component. In comparison of the syntax-related P600 to the irony-related P600, distributional differences were found. Moreover, for the P600 time window (i.e., 500–900 ms), different changes in theta power between the syntax and pragmatics effects were found, suggesting that different patterns of neural activity contributed to each respective effect. Thus, both late positivities seem to be differently sensitive to these two types of linguistic information, and might reflect distinct neurocognitive processes, such as reanalysis of the sentence structure versus pragmatic reanalysis.     

Framingham Risk Score with Cardiovascular Events in Chronic Kidney Disease

The Framingham Risk Score (FRS) was developed to predict coronary heart disease in various populations, and it tended to under-estimate the risk in chronic kidney disease (CKD) patients. Our objectives were to determine whether FRS was associated with cardiovascular events, and to evaluate the role of new risk markers and echocardiographic parameters when they were added to a FRS model. This study enrolled 439 CKD patients. The FRS is used to identify individuals categorically as “low” (<10% of 10-year risk), “intermediate” (10–20% risk) or “high” risk (≧ 20% risk). A significant improvement in model prediction was based on the −2 log likelihood ratio statistic and c-statistic. “High” risk (v.s. “low” risk) predicts cardiovascular events either without (hazard ratios [HR] 2.090, 95% confidence interval [CI] 1.144 to 3.818) or with adjustment for clinical, biochemical and echocardiographic parameters (HR 1.924, 95% CI 1.008 to 3.673). Besides, the addition of albumin, hemoglobin, estimated glomerular filtration rate, proteinuria, left atrial diameter >4.7 cm, left ventricular hypertrophy or left ventricular ejection fraction<50% to the FRS model significantly improves the predictive values for cardiovascular events. In CKD patients, “high” risk categorized by FRS predicts cardiovascular events. Novel biomarkers and echocardiographic parameters provide additional predictive values for cardiovascular events. Future study is needed to assess whether risk assessment enhanced by using these biomarkers and echocardiographic parameters might contribute to more effective prediction and better care for patients.     

It's funny because we think it's true: laughter is augmented by implicit preferences

This study tests the folk psychological belief that we find things funny because we think they are true. Specifically, it addresses the relationship between implicit preferences and laughter. Fifty-nine undergraduate Rutgers University students (33 females and 26 males) from ethnically diverse backgrounds were videotaped while watching a white stand-up comedian for 30 min. Positive emotional expression associated with laughter was later scored using the facial action coding system (FACS). Computer-timed Implicit Association Tests (IATs) were used to measure a subject's implicit preferences for traditional gender roles and racial preferences (blacks vs. whites). Results show that participants laughed more in response to jokes that matched their implicit preferences (e.g., those with stronger implicit preferences for whites laughed more at racially charged material). Implications for the evolution of humor, and laughter as a hard-to-fake signal of preferences, are discussed.     

Estimation of individual muscle force using elastography

Background

Estimation of an individual muscle force still remains one of the main challenges in biomechanics. In this way, the present study aimed: (1) to determine whether an elastography technique called Supersonic Shear Imaging (SSI) could be used to estimate muscle force, (2) to compare this estimation to that one provided by surface electromyography (EMG), and (3) to determine the effect of the pennation of muscle fibers on the accuracy of the estimation.

Methods and Results

Eleven subjects participated in two experimental sessions; one was devoted to the shear elastic modulus measurements and the other was devoted to the EMG recordings. Each session consisted in: (1) two smooth linear torque ramps from 0 to 60% and from 0 to 30% of maximal voluntary contraction, for the first dorsal interosseous and the abductor digiti minimi, respectively (referred to as “ramp contraction”); (2) two contractions done with the instruction to freely change the torque (referred to as “random changes contraction”). Multi-channel surface EMG recordings were obtained from a linear array of eight electrodes and the shear elastic modulus was measured using SSI. For ramp contractions, significant linear relationships were reported between EMG activity level and torque (R² = 0.949±0.036), and between shear elastic modulus and torque (R² = 0.982±0.013). SSI provided significant lower RMS_deviation between measured torque and estimated torque than EMG activity level for both types of contraction (1.4±0.7 vs. 2.8±1.4% of maximal voluntary contraction for “ramp contractions”, p<0.01; 4.5±2.3 vs. 7.9±5.9% of MVC for “random changes contractions”, p<0.05). No significant difference was reported between muscles.

Conclusion

The shear elastic modulus measured using SSI can provide a more accurate estimation of individual muscle force than surface EMG. In addition, pennation of muscle fibers does not influence the accuracy of the estimation.     

The voluntary control of heart rate under differential somatic restraint

The purpose of this investigation was to test the hypothesis that subjects(Ss) can exert voluntary, bidirectional heart rate(HR) control in the absence of concomitant changes in respiration rate(RR) and muscle activity(chin EMG). Feedback-assisted HR control was studied under four experimental conditions involving increasing levels of somatic restraint:(a) No somatic controls,(b) RR pacing,(c) EMG feedback, to maintain constant muscle activity, and(d) RR pacing plus EMG feedback. FifteenSs, eight females and seven males, were assigned to each of these four conditions(N=60). The HR control task involved ten increase and ten decrease trials in random order, each lasting 60 sec and preceded by a 60-sec resting baseline period. The results indicated thatSs produced significant HR increases \((\overline X \Delta = + 3.4bpm)\) , but were unable to effect decreases \((\overline X \Delta = - .3bpm)\) . Furthermore, the magnitude of the obtained HR changes did not differ by experimental condition, i.e., by level of somatic restraint. While concomitant bidirectional RR changes were found, the pacing procedure effectively reduced, and on HR increase trials, nearly eliminated, RR involvement. No increases in EMG activity were found on HR increase trials for any of the four experimental conditions, whereas all groups showed some EMG decreases on HR deceleration trials. With respect to the question of somatic mediation of autonomic control, it is concluded that HR increases produced under paced respiration “favor” an interpretation of unmediated HR acceleration.     

Brain activation by central command during actual and imagined handgrip under hypnosis.

The purpose was to compare patterns of brain activation during imagined handgrip exercise and identify cerebral cortical structures participating in "central" cardiovascular regulation. Subjects screened for hypnotizability, five with higher (HH) and four with lower hypnotizability (LH) scores, were tested under two conditions involving 3 min of 1) static handgrip exercise (HG) at 30% of maximal voluntary contraction (MVC) and 2) imagined HG (I-HG) at 30% MVC. Force (kg), forearm integrated electromyography, rating of perceived exertion, heart rate (HR), mean blood pressure (MBP), and differences in regional cerebral blood flow distributions were compared using an ANOVA. During HG, both groups showed similar increases in HR (+13 +/- 5 beats/min) and MBP (+17 +/- 3 mmHg) after 3 min. However, during I-HG, only the HH group showed increases in HR (+10 +/- 2 beats/min; P < 0.05) and MBP (+12 +/- 2 mmHg; P < 0.05). There were no significant increases or differences in force or integrated electromyographic activity between groups during I-HG. The rating of perceived exertion was significantly increased for the HH group during I-HG, but not for the LH group. In comparison of regional cerebral blood flow, the LH showed significantly lower activity in the anterior cingulate (-6 +/- 2%) and insular cortexes (-9 +/- 4%) during I-HG. These findings suggest that cardiovascular responses elicited during imagined exercise involve central activation of insular and anterior cingulate cortexes, independent of muscle afferent feedback; these structures appear to have key roles in the central modulation of cardiovascular responses.     

Hypnotic manipulation of effort sense during dynamic exercise: cardiovascular responses and brain activation.

The purpose of this investigation was to hypnotically manipulate effort sense during dynamic exercise and determine whether cerebral cortical structures previously implicated in the central modulation of cardiovascular responses were activated. Six healthy volunteers (4 women, 2 men) screened for high hypnotizability were studied on 3 separate days during constant-load exercise under three hypnotic conditions involving cycling on a 1) perceived level grade, 2) perceived downhill grade, and 3) perceived uphill grade. Ratings of perceived exertion (RPE), heart rate (HR), blood pressure (BP), and regional cerebral blood flow (rCBF) distributions for several sites were compared across conditions using an analysis of variance. The suggestion of downhill cycling decreased both the RPE [from 13 +/- 2 to 11 +/- 2 (SD) units; P < 0.05] and rCBF in the left insular cortex and anterior cingulate cortex, but it did not alter exercise HR or BP responses. Perceived uphill cycling elicited significant increases in RPE (from 13 +/- 2 to 14 +/- 1 units), HR (+16 beats/min), mean BP (+7 mmHg), right insular activation (+7.7 +/- 4%), and right thalamus activation (+9.2 +/- 5%). There were no differences in rCBF for leg sensorimotor regions across conditions. These findings show that an increase in effort sense during constant-load exercise can activate both insular and thalamic regions and elevate cardiovascular responses but that decreases in effort sense do not reduce cardiovascular responses below the level required to sustain metabolic needs.