Walking performance and aerobic and anaerobic metabolism of Carcinus maenas (L.) in sea water at 15°C

Walking performance of the shore crab Carcinus maenas (L.) in sea water at 15 °C was assessed. In large crabs there was an inverse relationship between fatigue time and speed; crabs ran for $\text{\$}\text{?}$10 min at 3.2 m·min^?1 and for only 2 min at 14 m·min^?1. There were linear relationships between oxygen consumption and walking speeds for small and large animals walking at up to 4 m·min^?1 Estimates of maximum oxygen consumption were proportional to W^0.13 whereas inactive consumption is proportional to W^0.44 this resulted in aerobic scope (i.e. the difference between inactive and maximal rates of oxygen consumption) remaining almost constant across a weight range of animals whereas the aerobic expansibility (maximal rates/inactive rates) declined from 7- to 4-fold with increasing size. After a 12-h period without handling (settled animals) the animals could immediately become active and reach maximal rates of oxygen consumption similar to those of animals handled 1 h before the experiment. The aerobic expansibility of these settled animals could range from 21 to 8 times their inactive rates of oxygen consumption in small and large animals respectively. After 10 min of exercise oxygen consumption and whole body lactate levels returned to pre-exercise values within 5 to 25 min. The net oxygen debts range from 16 to 64% of the net oxygen consumption increase during exercise in small and large animals respectively.Calculations of the energy gained from lactate accumulation indicated that the net aerobic energy production during walking was supplemented from 4 to 71 % by anaerobic metabolism in small and large animals respectively. With increasing animal size the decline in aerobic expansibility was offset by an increased capacity for lactate production so that the overall maximum energy production during sustained activity remained almost constant at around seven times the inactive rate. The cost of transport (the net increase in oxygen consumption per g per m) falls with increased walking speed and increased animal size.     

Comparative locomotor performance of marsupial and placental mammals

Marsupials are often considered inferior to placental mammals in a number of physiological characters. Because locomotor performance is presumed to be an important component of fitness, we compared marsupials and placentals with regard to both maximal running speeds and maximal aerobic speeds (=speed at which the maximal rate of oxygen consumption, VOlmax, is attained). Maximal aerobic speed is related to an animal's maximal sustainable speed, and hence is a useful comparative index of stamina.
Maximal running speeds of 11 species of Australian marsupials, eight species of Australian murid rodents, two species of American didelphid marsupials, and two species of American rodents were measured in the laboratory and compared with data compiled from the literature. Our values are greater than, or equivalent to, those reported previously. Marsupials and placentals do not differ in maximal running speeds (nor do Australian rodents differ from non-Australian rodents). Within these groups, however, species and families may differ considerably. Some of the interspecific variation in maximal running speeds is related to differences in habitat: species inhabiting open habitats (e.g. deserts) tend to be faster than are species from habitats with more cover, or arboreal species.
Maximal aerobic speeds (compiled from the literature) were higher in large species than in small species. However, marsupials and placentals show no general difference with regard to maximal aerobic speeds.
Maximal running speeds and maximal aerobic speeds for 18 species of mammals were not correlated, after correcting for correlations with body size. Thus, the fastest sprinters do not necessarily have high maximal aerobic speeds.     

Effects of Temperature and Water Loss on Terrestrial Locomotor Performance in Land Crabs: Integrating Laboratory and Field Studies

SYNOPSIS. Terrestrial and semi-terrestrial crustaceans are exposedto fluctuations in ambient temperature and conditions that favorevaporative water loss. These environmental stresses alter performancelimits in the laboratory and behavior in the field. The maximalrate of oxygen consumption, maximum aerobic speed, and endurancecapacity are greater at a body temperature (T_b) of 24°Cthan at 15°C or 30°C in the ghost crab, Ocypode quadrata.The total metabolic cost to move at the same relative speedis greater at a T_b of 24°C than at 15°C. Slower aerobickinetics at 15°C result in a smaller relative contributionof oxidative metabolism to total metabolic cost. However, therelative contributions from accelerated glycolysis are similarat both temperatures. When locomotion is intermittent, the totaldistance traveled before fatigue can be similar at Tbs of 15and 24°C but result from different movement and pause durationsat these temperatures. Performance limits of the ghost crabare negatively affected by dehydration and are sensitive torates of water loss. In the laboratory, endurance capacity ofthe fiddler crab, Uca pugilator, is greater at a T_b of 30°Cthan at 25°C. In the field, freely moving fiddler crabswith a T_b of 30°C travel at faster mean preferred speeds,as determined by motion analysis, than crabs at 25°C. Datafor land crabs support and advance general ectothermic modelsfor the effects of temperature and dehydration on locomotorperformance.     

Effects of age and physical activity status on the speed-aerobic demand relationship of walking.

Older adults tend to show lower preferred walking speeds and higher aerobic demands per distance walked than young adults. It has been suggested that a more sedentary life-style contributes to diminished musculoskeletal functioning, which in turn contributes to poorer economy of motion in the aged and sedentary adults. The purpose of this study was to quantify the speed-aerobic demand relationship during walking for old (greater than 65 yr of age) and young adults and to determine whether physical activity status affects this relationship. Aerobic demands for 30 young and 30 old individuals representing sedentary and physically active groups were measured as the subjects performed treadmill walking at seven speeds ranging from 0.67 to 2.01 m/s. All four age/physical activity groups displayed U-shaped speed-aerobic demand curves with minimum gross oxygen consumption per unit distance walked (ml.kg-1.km-1) at 1.34 m/s. A statistically significant age effect on walking aerobic demand was observed, with old subjects showing an 8% higher mean aerobic demand than the young subjects. This age-related effect was not associated with shifts in the speed at which aerobic demand was minimized or with the preferred walking speed of older individuals falling on a less economical portion of the speed-aerobic demand curve. Rather, it was speculated that declines in force-generating capacity of muscle in the aged may require recruitment of additional motor units and perhaps an additional proportion of less economical fast twitch muscle fibers to generate necessary forces. Physical activity status had no significant effect on walking aerobic demand.(ABSTRACT TRUNCATED AT 250 WORDS)     

Effects of obesity and sex on the energetic cost and preferred speed of walking.

The metabolic energy cost of walking is determined, to a large degree, by body mass, but it is not clear how body composition and mass distribution influence this cost. We tested the hypothesis that walking would be most expensive for obese women compared with obese men and normal-weight women and men. Furthermore, we hypothesized that for all groups, preferred walking speed would correspond to the speed that minimized the gross energy cost per distance. We measured body composition, maximal oxygen consumption, and preferred walking speed of 39 (19 class II obese, 20 normal weight) women and men. We also measured oxygen consumption and carbon dioxide production while the subjects walked on a level treadmill at six speeds (0.50-1.75 m/s). Both obesity and sex affected the net metabolic rate (W/kg) of walking. Net metabolic rates of obese subjects were only approximately 10% greater (per kg) than for normal-weight subjects, and net metabolic rates for women were approximately 10% greater than for men. The increase in net metabolic rate at faster walking speeds was greatest in obese women compared with the other groups. Preferred walking speed was not different across groups (1.42 m/s) and was near the speed that minimized gross energy cost per distance. Surprisingly, mass distribution (thigh mass/body mass) was not related to net metabolic rate, but body composition (% fat) was (r2= 0.43). Detailed biomechanical studies of walking are needed to investigate whether obese individuals adopt novel energy saving mechanisms during walking.     

The metabolic cost of changing walking speeds is significant,implies lower optimal speeds for shorter distances,and increases daily energy estimates

Humans do not generally walk at constant speed, except perhaps on a treadmill. Normal walking involves starting, stopping and changing speeds, in addition to roughly steady locomotion. Here, we measure the metabolic energy cost of walking when changing speed. Subjects (healthy adults) walked with oscillating speeds on a constant-speed treadmill, alternating between walking slower and faster than the treadmill belt, moving back and forth in the laboratory frame. The metabolic rate for oscillating-speed walking was significantly higher than that for constant-speed walking (6–20% cost increase for ±0.13–0.27 m s⁻¹ speed fluctuations). The metabolic rate increase was correlated with two models: a model based on kinetic energy fluctuations and an inverted pendulum walking model, optimized for oscillating-speed constraints. The cost of changing speeds may have behavioural implications: we predicted that the energy-optimal walking speed is lower for shorter distances. We measured preferred human walking speeds for different walking distances and found people preferred lower walking speeds for shorter distances as predicted. Further, analysing published daily walking-bout distributions, we estimate that the cost of changing speeds is 4–8% of daily walking energy budget.     

Optimal rate of work for mountaineers

The physiological responses of seven young male highlanders were recorded at high altitude while they were carrying loads (0, 25, 35, 45, and 55 kg) on snow at different speeds, supporting the loads on their backs by circular straps around the forehead. The rates of work calculated from the gross weight (body weight plus actual load in kg) multiplied by the speed of walking, m.min-1, ranged from 4,460 to 8,440 kg.m.min-1. The relationship between the rate of work and energy expenditure was rectilinear within the present range of values. The oxygen consumption (51.6 and 59.7 ml.min-1.kg-1 BW) for 55-kg load (at 4.09 and 4.64 km.h-1) possibly reached maximal aerobic capacity. At higher energy output at high altitude the subjects were exhausted after a short period of work. The proportion of increase of oxygen consumption per kg gross weight carried or per kg.m was almost constant up to a 55-kg experimental load. It is suggested that for day-to-day operations work should not be undertaken at more than 30-40% of maximal work capacity; a rate of work around 4,000 kg.m.min-1 (25-30 kg actual load at 3.0 to 3.5 km.h-1) may be considered as optimal for highlanders and porters at high altitude.     

Physiological responses to water-walking in middle aged women

The purpose of the present study was to examine the physiological responses to water-walking using the Flowmill, which has a treadmill at the base of a water-flume, in two groups of women. In the first group, the women were known to regularly swim and exercise in water (group A), while in the second, they did not routinely participate in water-exercise (group B). In both groups, twelve healthy female volunteers in their fifties participated in the study. All of the subjects walked in water using the Flowmill for the first time. Subjects completed four consecutive bouts of 4-minute duration at progressively increasing speeds (20, 30, 40, and 50 m.min-1), with 1-minute rests between each bout. In addition, water-velocity was adjusted to the walking speed of each bout. The water-depth of the Flowmill was the level of the xiphoid process. The water and room temperatures were 30.3 +/- 0.1 degrees C and 24.9 +/- 0.4 degrees C, respectively. In both groups, the relationship between walking speed and oxygen uptake (VO2) as well as that between walking speed and heart rate (HR) changed exponentially as the walking speed increased, and the relationship between HR and VO2 was linear. The relationship between HR and VO2 was similar in both groups, and there was no significant difference between the predicted maximal oxygen uptake (VO2max) of the two groups. VO2 and HR of group B during water-walking, however, were significantly higher than those of group A at all walking speeds. The results of this study clearly showed that experience in moving through the water strongly affects physiological responses to water-exercise, even when fitness levels are equivalent.     

Mesh selection and filtration efficiency of the Continuous Plankton Recorder

The filtration efficiency [volume filtered/(inlet aperture areax distance travelled through the water)] and retention of copepodsby the Continuous Plankton Recorder (CPR) were measured in seatrials. Filtration efficiency was independent of tow speed (n= 14 trials, range of speeds 5–13 knots, F_1.12 = 0.1,P = 0.73), but was influenced by the extent to which the bodyof the CPR was sealed. The retention of copepods on the silkfiltering mesh routinely used in CPRs did not differ significantlyfrom that predicted for a 270 µm nylon mesh and did notvary with tow speed.     

Capacity for sustained terrestrial locomotion in an insect: Energetics,thermal dependence,and kinematics

Summary The capacity for sustained, terrestrial locomotion in the cockroach. Blaberus discoidalis, was determined in relation to running speed, metabolic cost, aerobic capacity, and ambient temperature (T _a=15, 23, and 34°C; acclimation temperature=24°C). Steady-state thoracic temperature (T _tss) increased linearly with speed at each T _a.The difference between T _tss and T _awas similar at each experimental temperature with a maximum increase of 7°C. Steady-state oxygen consumption (VO_2ss) increased linearly with speed at each T _aand had a low thermal dependence (Q₁₀=1.0-1.4). The minimum cost of locomotion (the slope of the VO_2ss versus speed function) was independent of T _a.Cockroaches attained a maximal oxygen consumption (VO_2max). increased with T _afrom 2.1 ml O₂·g^-1·h^-1 at 15°C to 4.9 ml O₂·g^-1·h^-1 at 23°C, but showed no further increase at 34°C, VO_2max increased 23-fold over resting VO₂ at 23°C, 10-fold at 34°C, and 15-fold at 15°C. Endurance correlated with the speed at which VO_2max was attained (MAS, maximal aerobic speed). Temperature affected the kinematics of locomotion. compared to cockroaches running at the same speed, but higher temperatures (23–34°C), low temperature (15°C) increased protraction time, reduced stride frequency, and reduced stability by increasing body pitching. The thermal independence of the minimum cost of locomotion (C_min), the low thermal dependence of VO_2ss (i.e., y-intercept of the VO_2ss versus speed function), and a typical Q₁₀ of 2.0 for VO_2max combined to increase MAS and endurance in B. discoidalis when T _awas increased from 15 to 23°C. Exerciserelated endothermy enabled running cockroaches to attain a greater VO_2max, metabolic scope, and endurance capacity at 23°C than would be possible if T _tss remained equal to T _a. The MAS of B. discoidalis was similar to that of other arthropods that use trachea, but was 2-fold greater than ectotherms, such as salamanders, frogs, and crabs of a comparable body mass.Abbreviations T _a ambient temperature - T _t thoracic temperature - T _tss steady state thoracic temperature during exercise - T _trest thoracic temperature during rest - VO₂ oxygen consumption - VO_2rest oxygen consumption during rest - VO_2ss steady-state oxygen consumption during exercise - VO_2max maximal oxygen consumption; MAS maximum aerobic speed - C _min minimum cost of locomotion - t _end endurance time     

High-speed running performance: a new approach to assessment and prediction.

We hypothesized that all-out running speeds for efforts lasting from a few seconds to several minutes could be accurately predicted from two measurements: the maximum respective speeds supported by the anaerobic and aerobic powers of the runner. To evaluate our hypothesis, we recruited seven competitive runners of different event specialties and tested them during treadmill and overground running on level surfaces. The maximum speed supported by anaerobic power was determined from the fastest speed that subjects could attain for a burst of eight steps (approximately 3 s or less). The maximum speed supported by aerobic power, or the velocity at maximal oxygen uptake, was determined from a progressive, discontinuous treadmill test to failure. All-out running speeds for trials of 3-240 s were measured during 10-13 constant-speed treadmill runs to failure and 4 track runs at specified distances. Measured values of the maximum speeds supported by anaerobic and aerobic power, in conjunction with an exponential constant, allowed us to predict the speeds of all-out treadmill trials to within an average of 2.5% (R2 = 0.94; n = 84) and track trials to within 3.4% (R2 = 0.86; n = 28). An algorithm using this exponent and only two of the all-out treadmill runs to predict the remaining treadmill trials was nearly as accurate (average = 3.7%; R2 = 0.93; n = 77). We conclude that our technique 1) provides accurate predictions of high-speed running performance in trained runners and 2) offers a performance assessment alternative to existing tests of anaerobic power and capacity.     

Walking speed and area utilization of red king crab (<Emphasis Type="Italic">Paralithodes camtschaticus</Emphasis>) introduced to the Barents Sea coastal ecosystem

The red king crab (Paralithodes camtschaticus) was introduced in the Barents Sea in the 1960s and soon established a viable population. Proper management and exploitation of the Barents Sea king crab stock require better understanding of the spatial dynamics at different scales. This study examines the small-scale movement patterns of seven adult male crabs tracked for a period of up to one month from mid July to mid August at 150 m depth in a semi-enclosed fjord on the Russo-Norwegian border. The crabs were tagged with acoustic transmitters and their movements monitored with an acoustic positioning system. Low walking speeds (<0.01 m s⁻¹) were most frequent but the crabs could move at a maximum speed of 0.15 m s⁻¹ and walk an actual distance of up to 270 m over a period of one hour. However, the crabs usually moved within a relatively restricted area with mean hourly longest rectilinear distance varying from 26 to 64 m. The crabs alternated between periods of low and high activity, which could reflect feeding in and movements between food patches. The lack of a diel activity rhythm may be due to high light levels during the polar summer night, or a chemically mediated food search strategy.     

Intermittent locomotion increases endurance in a gecko

Nocturnal geckos can actively forage at low temperatures. A low minimum cost of locomotion allows greater sustainable speeds by partially offsetting the decrease in maximal oxygen consumption (VO2max) associated with low nocturnal temperatures. The nocturnality hypothesis (Autumn et al. 1997) proposes that the reduced cost of continuous locomotion is a shared, derived characteristic that increases the capacity to sustain locomotion at low temperatures. Yet many lizards move intermittently at speeds exceeding those that elicit VO2max. We exercised the frog-eyed gecko, Teratoscincus przewalskii, continuously and intermittently on a treadmill. At an exercise speed of 0.90 km h-1 (270% maximum aerobic speed), lizards alternating a 15-s exercise period with a 30-s pause period exhibited a 1.7-fold increase in distance capacity (total distance traveled before fatigue) compared with lizards exercised continuously at the same average speed (0.30 km h-1). The average aerobic cost of intermittent exercise was not significantly different from VO2max. Locomoting intermittently could augment the increase in endurance resulting from the low minimum cost of continuous locomotion in nocturnal geckos. Intermittent behavior could increase the endurance of lizard movement in general.     

Time Budgets, Thermoregulation, and Maximal Locomotor Performance: Are Reptiles Olympians or Boy Scouts?

SYNOPSIS. DO ectothermal vertebrates routinely make full useof their locomotor capacities in nature? We address this questionby asking whether reptiles ever sprint at maximum burst speedsand whether they often move at speeds near maximum aerobicallysustainable levels. Relevant data are largely anecdotal butsuggest that lizards (and perhaps other vertebrate ectotherms)do not routinely perform at maximal capacities. They appearto do so only in situations that have a critical impact on fitness.Nevertheless, active lizards do thermoregulate carefully suchthat they usually maintain the potential for performing at maximalcapacity. We consider alternative, but not exclusive, explanationsfor why reptiles might maintain apparently "excessive" capacitiesand conclude with suggestions for new field and laboratory studiesthat would more rigorously address these issues.     

Divergent selection for aerobic capacity in rats as a model for complex disease

Based upon ideas about evolution, we put forth the argumentthat the capacity to transfer energy via aerobic metabolismis such a central feature of mammalian biology, that it mustalso be the primary determinant of complex disease. From this,we hypothesized that artificial selection on low and high capacityfor aerobic exercise would create lines that can be used todefine the divide between health and disease. In 1996 we beganlarge-scale divergent selection for aerobic treadmill runningcapacity in a widely heterogeneous stock of rats (N:NIH). Byten generations we developed lines of low capacity runners (LCR)and high capacity runners (HCR) that on average differed by317%. As a correlated trait, body mass increased at each generationin the LCR while the body mass decreased in the HCR. The linesalso separated for key factors of systemic oxygen transportcapacity such as maximal oxygen consumption (VO₂max), tissueperfusion, capillary density, and oxidative enzyme activity(citrate synthase and B-HAD). We also tested our hypothesisthat differences in aerobic energy transfer would produce ratsthat contrast for risk factors associated with complex disease.Indeed, the lines separated for cardiovascular risk factorsincluding differences in blood pressure, cardiac contractility,visceral adiposity, plasma free fatty acids, and triglycerides.The decrease in aerobic capacity was also associated with lowamounts of several proteins required for mitochondrial function.     

Measurements of aerobic metabolism of a school of horse mackerel at different swimming speeds

Oxygen consumption rates were measured in a school of 56 horse mackerel Trachurus trachurus while at rest and while swimming at steady sustained speeds. Resting values of 38.76 and 42.10mg O₂ kg^?1 h^?1 were measured in a sealed cylindrical tank (535 l) while observing that the fish school remained neutrally buoyant and inactive with only gentle pectoral fin movements and no swimming motion. The same school was trained to swim with projected light patterns within a 10-m diameter annular doughnut respirometer. The oxygen consumption increased from the resting level through 51 mg O₂ kg^?1 h^?1 at the slowest swimming speeds of 0.29 m s^?1 (0.95 L s^?1) to around 259 mg O₂ kg^?1 h^?1 at the higher measured swimming speed of 0.87 m s^?1 (2.82 L s^?1). The data fitted a curve where oxygen consumption rose in proportion to velocity to the power of 2.56 with the intercept at the resting level. The maximum sustained speed (80 min) of 1.12 m s^?1 (3.63 Ls^?1) was not achieved within the respirometer but corresponded to an estimated oxygen consumption of 458.33 mg O₂ kg^?1 h^?1 giving a scope for aerobic activity of 419.02 mg O₂ kg^?1 h^?1. At a speed of 0.87 m s^?1, there was a lower bound on the aerobic efficiency of at least 38% and at 1.12 m s^?1, the highest aerobic speed, of 40%. Sustained speeds swum in a curved path as here should be increased by 5% for a straight path giving a maximum sustained 80 min speed of 1.18 m s^?1.     

Rush and grab strategies in foraging marine endotherms: the case for haste in penguins

The speed at which air-breathing marine predators that forage by diving should swim is likely to depend on a variety of factors that differ substantially from those relevant in animals for which access to oxygen is unlimited. We used loggers attached to free-living penguins to examine the speed at which three species swam during periods searching for prey and compared this to their speeds during actual prey pursuit. All penguin species appeared to travel at similar speeds around 2 m/s during normal commuting between the surface and feeding depths, which accords closely with minimum costs of transport. However, Adélie penguins, Pygoscelis adeliae, slowed down to feed, Magellanic penguins, Spheniscus magellanicus, speeded up and king penguins, Aptenodytes patagonicus, travelled at a variety of speeds, although mean speed did not change from normal commuting. Since energy expenditure, and therefore oxygen usage, in swimming animals increases with the cube of the speed, we hypothesized that prey escape speed (a function of prey size) and prey density would prove critical in determining optimum pursuit speeds in predators. Simple models of this type help explain why it is that some penguin species apparently benefit by increasing speed to capture prey while others benefit by decreasing speed.     

Skylark optimal flight speeds for flying nowhere and somewhere

Flight is energetically very costly. For birds the mechanicalpower in relation to airspeed is characterized by a U-shapedfunction. From this function we can derive optimal flight speedsassociated with minimum power (V_mp), minimum cost of transport(V_mr) and minimum overall time of migration (V_mt). Since flightis energetically so costly, aerial displays and song flightcan potentially serve as signals reliably indicating the individualquality or resource potential of the signaler. In order to maximizethe amount of song flight produced, we expect V_mp during songflight, while during migration we rather expect V_mr or V_mv Wecompared flight speeds of skylarks (Alauda arvensis) duringsong flight and migration flight, respectively. In this speciespredicted V_mp = 5.5 m/s, V_mr = 10.5 m/s, and V_mt = 12.1 m/s.The preferred airspeed during song flight did not differ significantlyfrom the predicted V_mp, while airspeed during migration wassignificantly higher than V_mr and Vmp indicating that flightspeed is a flexible trait that birds adjust to different situations.Why the skylarks speed up so much on migration is still unclear,but it may be that due to the shape of the predicted power curve,variation in cost of transport at high speeds is relativelysmall.     

Aerobic metabolic scope and swimming performance in juvenile cod, Gadus morhua L.

Juvenile cod (Gadus morhua) were made to swim in a tunnel respirometer to determine the oxygen consumption during swimming at different speeds. Results were compared with measurements of standard and active metabolic rates in static respirometers before and after intense exercise. The oxygen consumption at maximum sustainable swimming speed was considerably lower than the peak oxygen consumption following exhausting exercise. It is suggested that these fish have a poorly developed system of aerobic (red) locomotor muscles which do not normally make a major demand upon oxygen consumption. Apparent specific dynamic action following feeding and repayment of oxygen debt following anaerobic exercise can each give rise to greater rates of oxygen consumption. Following exhausting exercise there is a delay of about 1 h before oxygen consumption reaches a peak level some 40% higher than the peak level observed during sustained swimming.