Is the rate of whole-body nitrogen elimination influenced by exercise?

BACKGROUND: Because it has earlier been shown that exercise 24 or two hours pre-dive may suppress the appearance of venous gas bubbles (VGB) in connection with the dive, we studied whether exercise before or during N2 elimination would influence the rate of the latter. Nitrogen elimination was recorded in eight volunteers breathing a normoxic O2+argon mixture for two hours. The N2 washout was preceded two (Condition A) or 24 hours (Condition B) earlier, by one hour of exercise at 85% VO2max (two hours of exercise interspersed with two hours of rest). In separate experiments, exercise at -40% of VO2max was performed throughout the two-hour washout (Condition C), and control experiments (Condition D) with denitrogenation without exercise were also performed. RESULTS: There were no significant differences among conditions for the total N2 eliminated (904 +/- 196 mL). The half-times of N2 washout for A (35.2 +/- 10.8 minutes) and B (31.9 +/- 8.6 minutes) did not differ from control washouts. The rate of washout in C increased 14% compared to D (half-time: 30.4 +/- 7.6 vs. 34.5 +/- 7.8 minutes, p = 0.002), and correlated with cardiac output. CONCLUSION: Exercise 24 or two hours pre-N2 washout did not affect it, suggesting that the decreased VGB scores noted by others in dives preceded by conditions similar to A and B are not due to changes in nitrogen exchange but rather to factors related to bubble formation and/or appearance. That N2 elimination is enhanced by concomitant exercise makes physiological sense but does not necessarily explain the observation by others of a reduced risk of decompression sickness with exercise before diving.

Energy cost of breathing at depth: effect of respiratory muscle training.

INTRODUCTION: Respiratory muscle training against resistance (RRMT) increases respiratory muscle strength and endurance as well as underwater swimming endurance. We hypothesized that the latter is a result of RRMT reducing the high energy cost of breathing at depth. METHODS: Eight subjects breathed air in a hyperbaric chamber at 55 fsw, both before and after RRMT. They rested for 10 minutes, cycled on an ergometer for 10 minutes (100 W), rested for 10 minutes, and then, while still at rest, they voluntarily mimicked the breathing pattern recorded during the exercise (isocapnic simulated exercise ventilation, ISEV). RESULTS: Post-RRMT values of V(E) at rest, exercise and ISEV were not different from those recorded pre-RRMT. Pre-RRMT minute-ventilation (V(E)) during ISEV was not different from the exercise ventilation (49.98 +/- 10.41 vs. 47.74 +/- 8.44 L/minute). The end-tidal PCO2 during ISEV and exercise were not different (44.26 +/- 2.54 vs. 44.49 +/- 4.49 mmHg) or affected by RRMT. Oxygen uptake (VO2) was 0.32 +/- 0.08 L/ minute at rest, 1.78 +/- 0.15 during exercise pre-RRMT, and not different post-RRMT. During ISEV, VO2 decreased significantly from pre-RRMT to post-RRMT (0.46 +/- 0.06 vs. 0.36 +/- 0.11 L/minute). Post-RRMT delta VO2/delta V(E) was significantly lower during ISEV than pre-RRMT (0.0094 +/- 0.0021 L/L vs. 0.0074 +/- 0.0023 L/L). CONCLUSION: RRMT significantly reduced the energy cost of ventilation, measured as delta VO2/delta V(E) during ISEV, at a depth of 55 fsw. Whether this change was due to reduced work of breathing and/or increased efficiency of the respiratory muscles remains to be determined.

Energetics of swimming: a historical perspective.

The energy cost to swim a unit distance (C(sw)) is given by the ratio E/v where E is the net metabolic power and v is the swimming speed. The contribution of the aerobic and anaerobic energy sources to E in swimming competitions is independent of swimming style, gender or skill and depends essentially upon the duration of the exercise. C(sw) is essentially determined by the hydrodynamic resistance (W(d)): the higher W(d) the higher C(sw); and by the propelling efficiency (η(P)): the higher η(P) the lower C(sw). Hence, all factors influencing W(d) and/or η(P) result in proportional changes in C(sw). Maximal metabolic power E max and C(sw) are the main determinants of swimming performance; an improvement in a subject's best performance time can more easily be obtained by a reduction of C sw) rather than by an (equal) increase in E max (in either of its components, aerobic or anaerobic). These sentences, which constitute a significant contribution to today's knowledge about swimming energetics, are based on the studies that Professor Pietro Enrico di Prampero and his co-workers carried out since the 1970s. This paper is devoted to examine how this body of work helped to improve our understanding of this fascinating mode of locomotion.

Influence of exercise on nutritional requirements.

There is no consensus on the best diet for exercise, as many variables influence it. We propose an approach that is based on the total energy expenditure of exercise and the specific macro- and micronutrients used. di Prampero quantified the impact of intensity and duration on the energy cost of exercise. This can be used to determine the total energy needs and the balance of fats and carbohydrates (CHO). There are metabolic differences between sedentary and trained persons, thus the total energy intake to prevent overfeeding of sedentary persons and underfeeding athletes is important. During submaximal sustained exercise, fat oxidation (FO) plays an important role. This role is diminished and CHO's role increases as exercise intensity increases. At super-maximal exercise intensities, anaerobic glycolysis dominates. In the case of protein and micronutrients, specific recommendations are required. We propose that for submaximal exercise, the balance of CHO and fat favors fat for longer exercise and CHO for shorter exercise, while always maintaining the minimal requirements of each (CHO: 40% and fat: 30%). A case for higher protein (above 15%) as well as creatine supplementation for resistance exercise has been proposed. One may also consider increasing bicarbonate intake for exercise that relies on anaerobic glycolysis, whereas there appears to be little support for antioxidant supplementation. Insuring minimal levels of substrate will prevent exercise intolerance, while increasing some components may increase exercise tolerance.

The underwater environment: cardiopulmonary, thermal, and energetic demands.

Water covers over 75% of the earth, has a wide variety of depths and temperatures, and holds a great deal of the earth's resources. The challenges of the underwater environment are underappreciated and more short term compared with those of space travel. Immersion in water alters the cardio-endocrine-renal axis as there is an immediate translocation of blood to the heart and a slower autotransfusion of fluid from the cells to the vascular compartment. Both of these changes result in an increase in stroke volume and cardiac output. The stretch of the atrium and transient increase in blood pressure cause both endocrine and autonomic changes, which in the short term return plasma volume to control levels and decrease total peripheral resistance and thus regulate blood pressure. The reduced sympathetic nerve activity has effects on arteriolar resistance, resulting in hyperperfusion of some tissues, which for specific tissues is time dependent. The increased central blood volume results in increased pulmonary artery pressure and a decline in vital capacity. The effect of increased hydrostatic pressure due to the depth of submersion does not affect stroke volume; however, a bradycardia results in decreased cardiac output, which is further reduced during breath holding. Hydrostatic compression, however, leads to elastic loading of the chest wall and negative pressure breathing. The depth-dependent increased work of breathing leads to augmented respiratory muscle blood flow. The blood flow is increased to all lung zones with some improvement in the ventilation-perfusion relationship. The cardiac-renal responses are time dependent; however, the increased stroke volume and cardiac output are, during head-out immersion, sustained for at least hours. Changes in water temperature do not affect resting cardiac output; however, maximal cardiac output is reduced, as is peripheral blood flow, which results in reduced maximal exercise performance. In the cold, maximal cardiac output is reduced and skin and muscle are vasoconstricted, resulting in a further reduction in exercise capacity.

Tumbaga object from the early classic period, found at altun ha, british honduras (belize).

Excavations at Altun Ha, British Honduras (Belize), have yielded a claw-shaped bead of tumbaga, a gold-copper alloy, occurring as part of an Early Classic offering. Stratigraphic evidence plus radiocarbon dates place the cache at or before A.D. 500, and stylistic elements indicate a source for the bead in the Coclé culture of central Panama.

Ancient maya mercury.

Discovery of mercury in an ancient Maya offering at Lamanai, Belize, has stimulated examination of possible sources of the material in the Maya area. Two zones of cinnabar and native mercury deposits can be defined in the Maya highlands, and the presence of the native metal suggests that the ancient Maya collected rather than extracted the mercury from ore.

On the fringes of conquest: maya-spanish contact in colonial belize.

The defeat of the Aztecs of Mexico by Hernán Cortés in 1521 was but the beginning of a long and torturous conquest of Central America that did not always result in the mastery of people and production for which the Spanish had hoped. The Maya of the resource-poor Yucatán peninsula were spared the heavy colonial hand that held fast to central Mexico and its riches. In addition, the dense forests of the peninsula served as a haven for refugees fleeing oppressive conditions in colonial towns. Despite the paucity of documentary information on Maya communities of the frontier, knowledge of Maya-Spanish relations in the 16th and 17th centuries has advanced in recent years through archeological and ethnohistorical research. Work in one region of the Maya lowlands has brought us closer to an understanding of the early interaction of the rulers and the ruled.

Active and passive drag: the role of trunk incline.

The aim of this study was to investigate the role of trunk incline (TI) and projected frontal area (A(eff)) in determining drag during active/passive measurements. Active drag (D(a)) was measured in competitive swimmers at speeds from 0.6 to 1.4 m s(-1); speed specific drag (D(a)/v(2)) was found to decrease as a function of v (P < 0.001) to indicate that the human body becomes more streamlined with increasing speed. Indeed, both A(eff) and TI were found to decrease with v (P < 0.001) whereas C(d) (the drag coefficient) was found to be unaffected by v. These data suggest that speed specific drag depend essentially on A(eff). Additional data indicate that A(eff) is larger during front crawl swimming than during passive towing (0.4 vs. 0.24 m(2)). This suggest that D(a)/v(2) is larger than D(p)/v(2) and, at a given speed, that D(a) is larger than D(p).

Aerobic cost in elite female adolescent swimmers.

Maximal performance in swimming depends on metabolic power and the economy of swimming. Thus, the energy cost of swimming (economy= VO(2)/V, C(s)) and maximal aerobic power (VO(2max)) in elite young female swimmers (n=10, age: 15.3+/-1.5 years) and their relationships to race times (50-1,000 m) and national ranking were examined. VO(2) increased exponentially with velocity (V), (VO(2)=5.95+(-10.58 V)+5.84 V(2)) to a maximal VO(2) of 2.71+/-0.50 L x min(-1) (46.7+/-8.2 mL x kg(-1) x min(-1)) at a free swimming velocity of 1.37+/-0.07 m x s(-1). C(s) was constant up to 1.2 m x s(-1) (21.5 mL x m(-1)), however was significantly higher at 1.36 m x s(-1) (27.3 mL x m(-1)). Peak [La] was 5.34+/-2.26 mM. C(s) expressed as a percentage of Cs at maximal swimming velocity was significantly correlated with race times and ranking across a number of distances. The data for these elite females demonstrate that the energy cost of swimming is a good predictor of performance across a range of distances. However, as swimming performance is determined by a combination of factors, these findings warrant further examination.

Respiratory muscle training improves swimming endurance at depth.

Respiratory muscle training (RMT) has been shown to improve divers swimming endurance at 4 feet of depth; however, its effectiveness at greater depths, where gas density and the work of breathing are substantially elevated has not been studied. The purpose of this study was to examine the effects of resistance respiratory muscle training (RRMT) on respiratory function and swimming endurance at 55 feet of depth (270.5 kPa). Nine male subjects (25.9 +/- 6.8 years) performed RRMT for 30 min/day, 5 d/ wk, for 4 wks. Pre- and Post RRMT, subjects swam against a pre-determined load (70% VO2 max) until exhausted. As indices of respiratory muscle strength, maximal inspiratory and expiratory pressures were measured before and immediately following the swims pre- and post-RRMT. These measurements showed that ventilation was significantly lower during the swims and, at comparable swim duration, that the respiratory muscles were considerably less fatigued following RRMT. The reduced ventilation was due to a lower breathing frequency following RRMT. The ventilatory changes following RRMT coincided with significantly increased swimming time to exhaustion (approximately 60%, 31.3 +/- 11.6 vs. 49.9 +/- 16.0 min, pre- vs. post-RRMT, p < 0.05). These results suggest respiratory muscle fatigue limits swimming endurance at depth as well as at the surface and RRMT improves performance.

Statin therapy depresses total body fat oxidation in the absence of genetic limitations to fat oxidation.

Cholesterol lowering drugs are associated with myopathic side effects in 7% of those on therapy, which is reversible in most, but not all patients. This study tested the hypothesis that total body fat oxidation (TBFO) is reduced by statins in patients with genetic deficiencies in FO, determined by white blood cells (FOwbc) and by molecular analysis of common deficiencies, and would cause intolerance in some patients. Six patients on statin therapy without myopathic side effects (tolerant) and 7 patients who had previously developed statin-induced myopathic symptoms (intolerant) (age = 58 +/- 8.25 yrs, ht. = 169 +/- 11 cm, and wt. = 75.4 +/- 14.2 kg) were tested for TBFO (Respiratory Exchange Ratio, RER) pre- and during exercise. FOwbc was not significantly different between tolerant and intolerant (0.261 +/- 0.078 vs. 0.296 +/- 0.042 nmol/h per 10(9) wbc), or normals (0.27 +/- 0.09 nmol/h per 10(9) wbc) and no common molecular abnormalities were found. Pre-exercise RER (0.73 +/- 0.05 vs. 0.84 +/- 0.05) was significantly lower in the intolerant group and the VO2 at RER = 1.0 (1.27 +/- 0.32 vs. 1.87 +/- 0.60 L/min) greater than the tolerant. Post-exercise lactates were not different between groups. Although dietary fat intake was not different, blood lipoprotein levels, particularly triglycerides were 35% lower in tolerant than previously intolerant. TBFO and blood lipoproteins were reduced in tolerant patients in spite of the absence of genetic limitations, but not in the intolerant group as hypothesized. Although not conclusive, these data suggest the need for a prospective study of the effects of statins on fat oxidation.

Optimization of fin-swim training for SCUBA divers.

Underwater swimming is a unique exercise and its fitness is not accomplished by other types of training. This study compared high intensity intermittent fin-swim training (HIIT) with moderate intensity continuous (MICT). Divers (n = 20; age = 23 +/- 4 yrs; weight = 82.57 +/- 10.38 kg; height = 180 +/- 6 cm) were assigned to MICT (65%-75% heart rate max (HRmax), for 45 min) or HIIT three 10 min swims/rest cycles (77%, 83%, and 92% HRmax, respectively) for 50 min. They trained using snorkel and fins at the surface paced by an underwater light system 3 times per week for 4 weeks. Swim tests were the energy cost of swimming, VO2max and timed endurance swim (at 70%/VO2max). The VO2 was a non-significantly reduced at any velocity with either HIIT or MICT. Maximal swim velocity increased after HIIT (10%) (p < or = 0.05) but not after MICT (p > 0.05). VO2max increased 18% after HIIT and 6% after MICT (p < or = 0.05). The endurance times increased 131% after HIIT and 78% after MICT (p < or = 0.05), and in spite of this post-swim lactate was not significantly different and averaged 4.69 +/- 1.10mM (p > 0.05). Although both training methods significantly improved fin swimming performance with similar time commitments, the HIIT improved VO2max and endurance more than MICT (p < or = 0.05). As no improvements in ventilation were observed, combining HIIT with respiratory muscle training could optimize diver swim fitness.

Resistive respiratory muscle training improves and maintains endurance swimming performance in divers.

Respiratory work is increased during exercise under water and may lead to respiratory muscle fatigue, which in turn can compromise swimming endurance. Previous studies have shown that respiratory muscle training, conducted five days per week for four weeks, improved both respiratory and fin swimming endurance. This training (RRMT-5) consisted of intermittent vital capacity breaths (twice/minute) against spring loaded breathing valves imposing static and resistive loads generating average inspiratory pressures of approximately 40 cmH2O and expiratory pressures of approximately 47 cmH2O. The purpose of the present study (n = 20) was to determine if RRMT 3 days per week (RRMT-3) would give similar improvements, and if continuing RRMT 2 days per week (RRMT-M) would maintain the benefits of RRMT-3 in fit SCUBA divers. Pulmonary function, maximal inspiratory (P(insp)) and expiratory pressures (P(exp)), respiratory endurance (RET), and surface and underwater (4 fsw) fin swimming endurance were determined prior to and after RRMT, and monthly for 3 months. Pulmonary function did not significantly improve after either RRMT-3 or RMMT-5; while P(insp) (20 and 15%) and P(exp) (25 and 11%), RET (73 and 217%), surface (50 and 33%) and underwater (88 and 66%) swim times improved. VO2, VE and breathing frequency decreased during the underwater endurance swims after both RRMT-3 and RRMT-5. During RRMT-M P(insp) and P(exp) and RET and swimming times were maintained at post RRMT-3 levels. RRMT 3 or 5 days per week can be recommended to divers to improve both respiratory and fin swimming endurance, effects which can be maintained with RRMT twice weekly.

Effects of respiratory muscle training on respiratory CO2 sensitivity in SCUBA divers.

Typically, ventilation is tightly matched to CO2 production. However, in some cases CO2 is retained (SCUBA diving). One factor behind hypoventilation in divers may be low respiratory CO2 sensitivity. If this is due to inadequate respiratory muscle performance it might be remedied by respiratory muscle training (RMT). We retrospectively investigated respiratory CO2 sensitivity prior to and after RMT in several groups of SCUBA divers. CO2 sensitivity (slope of expired ventilation as a function of inspired PCO2) was measured with a rebreathing technique in 35 subjects with diving experience. RMT consisted of either isocapnic hyperventilation or intermittent vital capacity breaths (twice/minute) against spring loaded breathing valves imposing static and resistive loads generating average inspiratory pressures of approximately 40 cmH2O and expiratory pressures of approximately 47 cmH2O; RMT was performed 30 min/day, 3 or 5 days/week for 4 weeks. Based on pre-RMT CO2 sensitivity the subjects were divided into three groups: low sensitivity: < 2 l/min/mmHg PCO2, normal: 2-4 l/min/mmHg, and high sensitivity: > 4 l/min/mmHg of inspired PCO2. The normal group had a Pre-RMT CO2 sensitivity of 2.88 +/- 0.60 and a post RMT sensitivity of 2.51 +/- 0.88 l/min/mmHg (Mean +/- SD, n = 19, p = n.s). Response in low sensitivity subjects increased from 1.41 +/- 0.32 to 2.27 +/- 0.53 (n = 10, p = 0.002,) while in the high sensitivity group it decreased from 5.41 +/- 1.25 to 2.90 +/- 0.32 l/min/mmHg (n = 6, p = 0.003). These preliminary findings showed that 46% of the subjects had abnormal sensitivity, and suggest that RMT may normalize it in hypo- and hyper-ventilating divers. If the present results are verified, RMT may be an effective means of enhancing safety in CO2 retaining divers.

The influence of drag on human locomotion in water.

Propulsion in water requires a propulsive force to overcome drag. Male subjects were measured for cycle frequency, energy cost and drag (D) as a function of velocity (V), up to maximal V, for fin and front crawl swimming, kayaking and rowing. The locomotion with the largest propulsive arms and longest hulls traveled the greatest distance per cycle (d/c) and reached higher maximal V. D while locomotoring increased as a function of V, with lower levels for kayaking and rowing at lower Vs. For Vs below 1 m/s, pressure D dominated, while friction D dominated up to 3 m/s, after which wave D dominated total D. Sport training reduced the D, increased d/c, and thus lowered C and increased maximal V. Maximal powers and responses to training were similar in all types of locomotion. To minimize C or maximize V, D has to be minimized by tailoring D type (friction, pressure or wave) to the form of locomotion and velocity.

Morphologic and functional alterations in aging rat muscle.

To understand better the causes of reduced contractile force in aging skeletal muscle, we performed a physiologic and morphologic analysis of plantaris muscle in old rats. The peak twitch tension (Fmax) and rates of force development and relaxation were significantly lower in old (24 months old) rats than in young (six month old) rats. In teased muscle fiber preparations, there was a 5% reduction in the mean number of fibers in the aging plantaris muscle. Histologically, a net loss of fibers occurred only in the muscle belly. Histochemically, fewer Type I fibers were seen in the belly and proximal regions, whereas distally fewer Type IIa fibers were seen. The loss of Types I and IIa oxidative fibers suggested a conservation of fast-twitch Type IIb fibers in a fast-twitch muscle. The relatively small loss of muscle fibers does not explain the large decline in muscle contractile performance which, despite established doctrine, was independent of muscle mass, fiber number or size, or number of fast-twitch fibers. The reduced force production in aging rat muscle appears to be due to a defect in excitation, contraction performance or metabolic activity, rather than a purely anatomical abnormality of muscle.

Blood flow distribution in dog gastrocnemius muscle at rest and during stimulation.

The distribution of blood flow within the isolated perfused dog gastrocnemius muscle (weight 100-240 g) was studied by intra-arterial injection of radioactively labeled microspheres (diameter 15 micron) at rest and during supramaximal stimulation to rhythmic isotonic tetanic contractions of varied frequency against varied loads. After the experiment the muscle was cut into 180-250 pieces of approximately 0.75 g each, and the blood flow to each muscle piece was determined from its radioactivity. The inhomogeneity of blood flow was represented as the frequency distribution of the ratios of regional specific blood flow, i.e., blood flow per unit tissue weight of the piece, QR, to the overall specific blood flow of the muscle, Q. The QR/Q values for the individual pieces of a muscle were found to vary widely both at rest and during stimulation. With rising work load the frequency distribution had a tendency to broaden and flatten, indicating increasing perfusion inhomogeneity. On the average of the experiments, there was no significant difference in specific blood flow between the three anatomic components of the gastrocnemius (lateral and medial heads of gastrocnemius and flexor digitorum superficialis) nor between the superficial and deep portions within these anatomic components, only the distal third of the muscle was relatively less perfused compared with the proximal two-thirds. The considerable inhomogeneity of blood flow as revealed by microsphere embolization and by other methods is expected to exert important limiting effects on local O2 supply, particularly during exercise. Its neglect would lead to serious errors in the analysis of O2 supply to muscle tissue.

Fluid conservation in athletes: responses to water intake, supine posture, and immersion.

The roles of antidiuretic hormone (ADH) and aldosterone in the elicited diuretic responses of trained and untrained men to seated, supine, and head-out water immersed conditions were studied. Volunteers were comprised of groups of six untrained individuals, six trained swimmers, and six trained runners. Each subject underwent three protocols, six hours in a seated position, supine position, or immersion (35 degrees C water). The last two protocols were preceded and followed by 1 h of seated position. After 10 h of fasting, 0.5% body wt of water was drunk. One hour later the trained groups had higher urine osmolalities (P less than 0.05) and urinary excretion rates of ADH (P less than 0.05) and lower urine flow rates (P less than 0.05) than untrained subjects. Throughout the sitting protocol, urinary ADH was also higher in both trained groups (P less than 0.05). Both supine posture and immersion resulted in significant decreases in urinary ADH in the untrained subjects (P less than 0.05) but no changes wer noted in swimmers and only during the second hour of immersion in the runners (P less than 0.05). The natriuresis and kaliuresis were greater during immersion than in the supine position but plasma renin activity, measured only in trained groups, and plasma aldosterone, measured in the untrained group, were decreased similarly with both protocols. The increases in urinary sodium excretion and urine flow rate were lower in trained than untrained subjects during the supine and immersion protocols (P less than 0.05). The data are compatible with an increased osmotic but decreased volume sensitivity of ADH control in trained men.

Cardiovascular response to submaximal exercise in sustained microgravity.

Cardiac output (Q), heart rate (HR), blood pressure, and oxygen consumption (VO2) were measured repeatedly both at rest and at two levels of exercise in six subjects during microgravity exposure. Exercise was at 30 and 60% of the workload producing the individual's maximal VO2 in 1 G. Three of the subjects were on a 9-day flight, Spacelab Life Sciences-1, and three were on a 15-day flight, Spacelab Life Sciences-2. We found no temporal differences during the flights. Thus we have combined all microgravity measurements to compare in-flight values with erect or supine control values. At rest, Q in flight was 126% of Q erect (P < 0.01) but was not different from Q supine, and HR in flight was 81% of HR erect (P < 0.01) and 91% of HR supine (P < 0.05). Thus resting stroke volume (SV) in flight was 155% of SV erect (P < 0.01) and 109% SV supine (P < 0.05). Resting mean arterial blood pressure and diastolic pressure were lower in flight than erect (P < 0.05). Exercise values were considered as functions of VO2. The increase in Q with VO2 in flight was less than that at 1 G (slope 3.5 vs. 6.1 x min-1.l-1.min-1). SV in flight fell with increasing VO2, whereas SV erect rose and SV supine remained constant. The blood pressure response to exercise was not different in flight from erect or supine. We conclude that true microgravity causes a cardiovascular response different from that seen during any of its putative simulations.