Search

Evaluating fish feeding behaviours is imperative to understanding prey resource use in the field. Previous work on fish feeding has taken place almost exclusively in a laboratory setting, which may impose artificial restrictions. Thus, we aimed to evaluate the constraints the captive setting places on fish feeding behaviours. We recorded polymorphic (littoral and pelagic) bluegill sunfish (Lepomis macrochirus) feeding in the laboratory and the field using a high‐speed camera and underwater cameras. Following successful strikes video events, were digitised using ImageJ (laboratory) and Argus (field). Gape velocity, ram velocity, and body deceleration were higher in the field than in captive fish. Significantly greater gape velocity in field fish suggests that these fish feed with greater suction pressure than captive fish. Prey effects were detected, as brine shrimp feeding events were characterised by slower gape and ram velocities and a smaller gape. Feeding events on brine shrimp in the field were similar to feeding events on worms in the laboratory suggesting an artefact of training. These results indicate that feeding behaviours measured in the laboratory may not be representative of feeding behaviours in the wild. Further consideration of organismal performance and laboratory constraints should be taken in future functional studies.

A variety of fish species show habitat-related variation in traits associated with swimming performance and foraging behavior. This commonly manifests as a distinction between open water and shallow water littoral ecotypes. In bluegill sunfish (Lepomis macrochirus), open water fish exhibit greater energy economy and speed during sustained locomotion than those from the littoral, whereas littoral fish were more maneuverable than their open water counterparts. These distinctions are associated with variation in diet and foraging behavior and may represent a resource polyphenism that enhances fitness through more effective exploitation of particular habitat types. A lack of field data means that polyphenisms have not been placed in context with swimming behavior in the field. We have used 3D videography to quantify bluegill field swimming performance in open water and littoral habitats. This revealed patterns of performance variation that parallel the trait variation previously established in the laboratory. Open water fish utilized faster average swimming speeds than inshore fish, while indicators of nonlinearity and unsteadiness were greater in the littoral fish. There are, however, differences in propulsive behavior between the field and laboratory. Pectoral-fin-powered, median-paired fin swimming is rarely employed by open water fish. Field body-caudal fin swimming involves short sequences of propulsive tail beats interspersed with gliding, rather than the repeated propulsive cycles employed under steady-state conditions. This suggests a need to re-evaluate the applicability of steady-state performance traits to behavior and fitness in the field and highlights the general importance of obtaining field performance data.

Skeletal muscle has diverse mechanical roles during locomotion. In swimming fish, power-producing muscles work in concert with the accessory muscles of the fins which augment and control power transfer to the water. Although fin muscles represent a significant proportion of the locomotor muscle mass, their physiological properties are poorly characterized. To examine the relationship between muscle metabolism and the differing mechanical demands placed on distinct muscle groups, we quantified the aerobic and glycolytic capacities of the myotomal, pectoral and caudal muscles of bluegill sunfish. These were indicated by the activities of citrate synthase and lactate dehydrogenase, rate-limiting enzymes for aerobic respiration and glycolysis, respectively. The well-established roles of slow and fast myotomal muscle types in sustained and transient propulsive movements allows their use as benchmarks to which other muscles can be compared to assess their function. Slow myotomal muscle had the highest CS activity, consistent with meeting the high metabolic and mechanical power demands of body-caudal fin (BCF) swimming at the upper end of the aerobically supported speed range. The largest pectoral adductors and abductors had CS activities lower than the slow myotomal muscle, in line with their role supplying thrust for low-speed, low-power swimming. The metabolic capacities of the caudal muscles were surprisingly low and inconsistent with their activity during steady-state BCF swimming at high speeds. This may reflect adaptation to the observed swimming behavior in the field, which typically involved short bouts of BCF-propulsive cycles rather than sustained propulsive activity.

The drag coefficient bluegill sunfish Lepomis macrochirus was estimated from coasting deceleration as (mean ± SD) 0.0154 ± 0.0070 at a Reynolds number of 41,000 ± 14,000. This was within the coasting range in other species and lower than values obtained from dead drag measurements in this species and others. Low momentum losses during coasting may allow its use during intermittent propulsion to modulate power output or maximize energy economy.

Mobility is essential to the fitness of many animals, and the costs of locomotion can dominate daily energy budgets. Locomotor costs are determined by the physiological demands of sustaining mechanical performance, yet performance is poorly understood for most animals in the field, particularly aquatic organisms. We have used 3-D underwater videography to quantify the swimming trajectories and propulsive modes of bluegills sunfish (Lepomis macrochirus, Rafinesque) in the field with high spatial (1-3 mm per pixel) and temporal (60 Hz frame rate) resolution. Although field swimming trajectories were variable and nonlinear in comparison to quasi steady-state swimming in recirculating flumes, they were much less unsteady than the volitional swimming behaviors that underlie existing predictive models of field swimming cost. Performance analyses suggested that speed and path curvature data could be used to derive reasonable estimates of locomotor cost that fit within measured capacities for sustainable activity. The distinct differences between field swimming behavior and performance measures obtained under steady-state laboratory conditions suggest that field observations are essential for informing approaches to quantifying locomotor performance in the laboratory.

Locomotion is energetically expensive. This may create selection pressures that favor economical locomotor strategies, such as the adoption of low-cost speeds and efficient propulsive movements. For swimming fish, the energy expended to travel a unit distance, or cost of transport (COT), has a U-shaped relationship to speed. The relationship between propulsive kinematics and speed, summarized by the Strouhal number (St=fA/U, where f is tail beat frequency, A is tail tip amplitude in m and U is swimming speed in m s−1), allows for maximal propulsive efficiency where 0.2

Bluegill sunfish use intermittent propulsion during volitional swimming. The function of this propulsive mode during routine swimming has not been well quantified. At low speeds, propulsive cycle frequencies and amplitudes were constant, and average speed and power output were controlled by modulating coasting duration. This fixed-gear gait may accommodate muscle level constraints on power production. At higher speeds bluegills switched to a mixed power-modulation strategy, increasing speed and power through increased propulsive cycle frequency and reduced coasting time.

Bluegill sunfish Lepomis macrochirus exhibit intraspecific variation in their morphology and swimming performance based on habitat. The pelagic form has a relatively streamlined, fusiform body shape associated with greater steady-state swimming speed and energy economy. In contrast, littoral bluegill have deeper bodies with fins located farther from their center of mass to enhance maneuverability among littoral vegetation. Deeper body shapes have been associated with increased fast-start performance to escape predators or capture prey. We hypothesized that littoral bluegill, which have a deeper body shape, would exhibit greater fast-start performance than pelagic bluegill. A total of 29 bluegill (16 littoral, 13 pelagic) were caught by hook and line, and their fast-start performance was analyzed from high-speed video recordings. Body shape appears to be a poor predictor of fast-start performance. Contrary to our expectations, pelagic bluegill had a significantly higher peak velocity, peak acceleration, and angular velocity compared to littoral bluegill. Pelagic bluegill living among larger predators and foraging on mobile prey may be exposed to selection pressures that favor increased fast-start performance. Integrated studies of internal morphology and physiology are needed to fully understand the relationship between morphology and performance in this population.

Medicinal leeches (Hirudo verbana) thermoregulate with respect to their sanguivorous feeding behavior. Immediate postprandial preferences are for warmer than their initial acclimation temperature (Ta, 21 °C, Petersen et al. 2011), while unfed leeches have a lower preferred temperature (Tpref, 12.5 °C). This may reduce energy expenditure and defer starvation if feeding opportunities are limited. Energetic benefits may have an associated cost if low temperatures reduce mobility and the ability to locate further hosts. These costs could be limited if mobility is unimpaired at low temperatures, or if acclimation can restore locomotor performance to the levels at Ta. The transition from Ta to the unfed Tpref significantly reduced speed and propulsive cycle frequency during swimming, and extension and retraction rates during crawling. Aerobic metabolic rate was also reduced from 0.20±0.03 W kg−1 at Ta to 0.10±0.03 W kg−1 at Tpref. The Q10 values of 1.7–2.9 for energetic and swimming parameters indicate a substantial temperature effect, although part of the decline in swimming performance can be attributed to temperature-related changes in water viscosity. 6 weeks at Ta resulted in no detectable acclimation in locomotor performance or aerobic metabolism. The energetic savings associated with a lower Tpref in unfed leeches effectively doubled the estimated time until depletion of energy reserves. Given that some mobility is still retained at Tpref, and that acclimation is in itself costly, the energetic benefits of selecting cooler temperatures between feedings may outweigh the costs associated with reduced locomotor performance.

Escape maneuvers are essential to the survival and fitness of many animals. Escapes are frequently initiated when an animal is already in motion. This may introduce constraints that alter the escape performance. In fish, escape maneuvers and steady, body caudal fin (BCF) swimming are driven by distinct patterns of curvature of the body axis. Pre-existing muscle activity may therefore delay or diminish a response. To quantify the performance consequences of escaping in flow, escape behavior was examined in bluegill sunfish (Lepomis macrochirus) in both still-water and during steady swimming. Escapes executed during swimming were kinematically less variable than those made in still-water. Swimming escapes also had increased response latencies and lower peak velocities and accelerations than those made in still-water. Performance was also lower for escapes made up rather than down-stream, and a preference for down-stream escapes may be associated with maximizing performance. The constraints imposed by pre-existing motion and flow, therefore, have the potential to shape predator–prey interactions under field conditions by shifting the optimal strategies for both predators and prey.

SUMMARY We examined the hypothesis that structural features of the iliotibialis lateralis pars postacetabularis (ILPO) in guinea fowl allow this large muscle to maintain equivalent function along its anterior–posterior axis. The ILPO, the largest muscle in the hindlimb of the guinea fowl, is a hip and knee extensor. The fascicles of the ILPO originate across a broad region of the ilium and ischium posterior to the hip. Its long posterior fascicles span the length of the thigh and insert directly on the patellar tendon complex. However, its anterior fascicles are shorter and insert on a narrow aponeurosis that forms a tendinous band along the anterior edge of the muscle and is connected distally to the patellar tendon. The biarticular ILPO is actively lengthened and then actively shortened during stance. The moment arm of the fascicles at the hip increases along the anterior to posterior axis, whereas the moment arm at the knee is constant for all fascicles. Using electromyography and sonomicrometry, we examined the activity and strain of posterior and anterior fascicles of the ILPO. The activation was not significantly different in the anterior and posterior fascicles. Although we found significant differences in active lengthening and shortening strain between the anterior and posterior fascicles, the differences were small. The majority of shortening strain is caused by hip extension and the inverse relationship between hip moment arm and fascicle length along the anterior–posterior axis was found to have a major role in ensuring similar shortening strain. However, because the knee moment arm is the same for all fascicles, knee flexion in early stance was predicted to produce much larger lengthening strains in the short anterior fascicles than our measured values at this location. We propose that active lengthening of the anterior fascicles was lower than predicted because the aponeurotic tendon of insertion of the anterior fascicles was stretched and only a portion of the lengthening had to be accommodated by the active muscle fascicles.

Phenotypic plasticity in response to environmental cues can create distinct morphological types within populations. This variation in form, and potentially function, may be a factor in initiating population divergence and the formation of new species. Here we show the translation of sympatric, habitat-specific morphological divergence into performance differences in energy economy, maneuverability and steady-state locomotion. Littoral and pelagic bluegill sunfish ecomorphs show differences in performance that appear adaptive within their respective habitats: greater maneuverability in the heavily vegetated littoral; greater steady-state swimming speed and economy in the open-water pelagic. This represents a trade-off in unsteady versus steady swimming performance, likely because morphological features associated with maximizing maneuverability are incompatible with enhancing steady-swimming performance. This may constrain the direction of adaptive change, maintaining the divergence created by phenotypic plasticity. The combination of habitat specific sympatric adaptation and constraints imposed by performance trade-offs may be an important factor underlying the high rate of speciation in freshwater fishes from post-glacial lakes.

The body wall muscles of sanguivorous leeches power mechanically diverse behaviours: suction feeding, crawling and swimming. These require longitudinal muscle to exert force over an extremely large length range, from 145 to 46 per cent of the mean segmental swimming length. Previous data, however, suggest that leech body wall muscle has limited capacity for force production when elongated. Serotonin (5-HT) alters the passive properties of the body wall and stimulates feeding. We hypothesized that 5-HT may also have a role in allowing force production in elongated muscle by changing the shape of the length–tension relationship (LTR). LTRs were measured from longitudinal muscle stripsin vitroin physiological saline with and without the presence of 10 µM 5-HT. The LTR was much broader than previously measured for leech muscle. Rather than shifting the LTR, 5-HT reduced passive muscle tonus and increased active stress at all lengths. In addition to modulating leech behaviour and passive mechanical properties, 5-HT probably enhances muscle force and work production during locomotion and feeding.

Bluegill Lepomis macrochirus showed intraspecific morphological and behavioural differences dependent on the environment. Pelagic L. macrochirus had more fusiform bodies, a higher pectoral fin aspect ratio, a larger spiny dorsal fin area and pectoral fins located farther from the centre of mass than littoral L. macrochirus (P < 0·05). The shape of the body and pectoral fins, in particular, were suggestive of adaptation for sustained high-speed and economical labriform swimming. Littoral L. macrochirus had a deeper and wider body, deeper caudal fins and wider mouths than pelagic L. macrochirus (P < 0·05). Additionally, the soft dorsal, pelvic, anal and caudal fins of littoral L. macrochirus were positioned farther from the centre of mass (P < 0·05). The size and placement of these fins suggested that they will be effective in creating turning moments to facilitate manoeuvring in the macrophyte-dense littoral habitat.

SUMMARY Although mechanical linkages between the proximal and distal limb are present in a range of species, their functional significance is unknown. We have investigated the mechanical function of the flexor cruris lateralis pars pelvica (FCLP), flexor cruris lateralis pars accessoria (FCLA) and gastrocnemius intermedia (GI), a system of linked muscles spanning proximal and distal limb segments in the guinea fowl (Numida meleagris) hind limb. The FCLP, which is in the anatomical position of a hamstring muscle, is the primary component of the linkage. It is connected to the distal femur via the FCLA, the tarsometatarsus via the tendon of insertion of the GI and the common Achilles tendon, and the tibiotarsus via a distal tendon of insertion. The FCLP may, therefore, potentially exert moments at the hip, knee and ankle joints depending on the joint angles and the relative states of activation in the three muscles. Evidence presented here suggests that the GI and FCLA act as actively controlled links that alter distal action of the FCLP. The FCLP and GI are coactive in the late swing and early stance phases of the stride, forming a triarticular complex, and likely act together to resist and control ankle flexion immediately after foot-down in addition to providing hip extension and knee flexion moments. The FCLP and FCLA are coactive from mid-through to late stance, acting together as a uniarticular hip extensor. Available evidence suggests that this role of the FCLP and FCLA is of increased importance in inclined running and accelerations. This linkage between a proximal muscle and alternate distal connections allows for functional flexibility, both in terms of the site at which the muscle exerts force and the nature of the muscle's mechanical function. The interactions generated between the proximal and distal limb by linkages of this type suggest that less emphasis should be placed on the distinct functional roles of specific anatomical classes of muscle within proximal and distal limb segments.

The medicinal leech is a useful animal model for investigating undulatory swimming in the classroom. Unlike many swimming organisms, its swimming performance can be quantified without specialized equipment. A large blood meal alters swimming behavior in a way that can be used to generate a discussion of the hydrodynamics of swimming, muscle mechanics, hydrostatic skeletons, and the physiological features that allow leeches to deal with the volume increase and osmotic load imposed by the meal. Analyses can be carried out at a range of levels tailored to suit a particular class.

The locomotor system of sanguivorous leeches is presented with a unique challenge: how to maintain mobility while coping with a >500% increase in body mass during feeding. A meal of this size is likely to disrupt the function of the muscular hydrostat during swimming, reducing speed and increasing predation risks. We quantified the effects of feeding to satiety on swimming kinematics, and the time course of recovery of swimming performance post-feeding in the medicinal leech Hirudo verbana. There was a 5.07 ± 0.04-fold increase in mass during feeding (mean ±sem, n=7). Despite this, leeches were able to swim immediately after feeding, reaching 27% of their pre-feeding speed. Reduced speed was a consequence of a reduction in both swimming cycle frequency and stride length to 69 and 42% of the pre-feeding values, respectively. Recovery of swimming ability was rapid, despite a prolonged increase in body mass. Fifty per cent restoration of swimming speed was achieved in c. 1 h while body mass was still 4.2-fold greater than before feeding. Rapid mass and volume reduction immediately post-feeding, and the properties of the obliquely striated swimming muscles appear to aid recovery of swimming performance. Such features that aid post-feeding recovery of mobility may have been important in the evolution of leech sanguivory.

SUMMARY Many fish change gait within their aerobically supported range of swimming speeds. The effects of acute temperature change on this type of locomotor behavior are poorly understood. Bluegill sunfish swim in the labriform mode at low speeds and switch to undulatory swimming as their swimming speed increases. Maximum aerobic swimming speed ( U max ), labriform-undulatory gait transition speed ( U trans ) and the relationships between fin beat frequency and speed were measured at 14, 18, 22, 26 and 30°C in bluegill acclimated to 22°C. At temperatures below the acclimation temperature ( T a ), U max , U trans and the caudal and pectoral fin beat frequencies at these speeds were reduced relative to the acclimation level. At temperatures above T a there was no change in these variables relative to the acclimation level. Supplementation of oxygen levels at 30°C had no effect on swimming performance. The mechanical power output of the abductor superficialis, a pectoral fin abductor muscle, was measured in vitro at the same temperatures used for the swimming experiments. At and below T a , maximal power output was produced at a cycle frequency approximately matching the in vivo pectoral fin beat frequency. At temperatures above T a muscle power output and cycle frequency could be increased above the in vivo levels at U trans . Our data suggest that the factors triggering the labriform–undulatory gait transition change with temperature. Muscle mechanical performance limited labriform swimming speed at T a and below, but other mechanical or energetic factors limited labriform swimming speed at temperatures above T a .

Morphology, size and physiological properties change markedly across fish ontogeny. This impacts locomotor performance and organismal fitness, although the effects are unpredictable due to the complexity of phenotype–function relationships. Morphological and behavioral changes with growth are often paralleled by changes in habitat use, diet and vulnerability to predators. Our goal was to quantify the changes in external morphology and escape performance throughout post-larval development in bluegill sunfish ( Lepomis macrochirus ), and place these changes in context with known changes in habitat use in the field. Development into adult ecomorphs is associated with phenotypic plasticity in response to habitat-specific differences in diet. On this basis, we hypothesized that variation in morphology and performance would increase during bluegill ontogeny as diversification of adult ecomorphs occurred. However, we found that variation in phenotype and escape performance decreased during early ontogeny. Phenotypic variation expanded later in development, after fish gained access to the variety of habitats and food types that may favor phenotypic plasticity. Performance is predicted to decline with growth due to the differential scaling of inertia and cross-sectional area, a major determinant of muscle force. In contrast, acceleration increased with size, and velocity and acceleration increased more rapidly with size than predicted. Post-larval maturation in bluegill featured a shift to a deeper body shape, and an increase in the relative size of the anal and caudal fins. This was a likely factor in the deviation of escape performance scaling relationships from predictions based on geometric similarity.

Successful predator evasion is essential to the fitness of many animals. Variation in escape behaviour may be adaptive as it reduces predictability, enhancing escape success. High escape velocities and accelerations also increase escape success, but biomechanical factors likely constrain the behavioural range over which performance can be maximized. There may therefore be a trade-off between variation and performance during escape responses. We have used bluegill sunfish (Lepomis macrochirus) escape responses to examine this potential trade-off, determining the full repertoire of escape behaviour for individual bluegill sunfish and linking this to performance as indicated by escape velocity and acceleration. Fish escapes involve an initial C-bend of the body axis, followed by variable steering movements. These generate thrust and establish the escape direction. Directional changes during the initial C-bend were less variable than the final escape angle, and the most frequent directions were associated with high escape velocity. Significant inter-individual differences in escape angles magnified the overall variation, maintaining unpredictability from a predator perspective. Steering in the latter stages of the escape to establish the final escape trajectory also affected performance, with turns away from the stimulus associated with reduced velocity. This suggests that modulation of escape behaviour by steering may also have an associated performance cost. This has important implications for understanding the scope and control of intra- and inter-individual variation in escape behaviour and the associated costs and benefits.

SUMMARYLinking the mechanics and energetics of locomotion in vertebrates has been hampered by a lack of information regarding the energy use of individual skeletal muscles in vivo. Here, we present a review of the available data concerning the relationship between the rates of skeletal muscle blood flow and oxygen consumption(V̇O2). In active muscle, during aerobically supported exercise, there is a linear relationship between these variables, irrespective of the muscle fiber type and intensity of exercise through most of the aerobic exercise range. We conclude that the rate of blood flow is the best available indicator of aerobic metabolic rate in multiple individual muscles or regions of muscles during locomotion. The practical considerations of using the injectable microsphere technique to measure muscle blood flow in this context are discussed.

SUMMARYWe examined the changes in muscle energy use in guinea fowl running at 1.5 m s-1 either unloaded, or carrying trunk loads equal to 23% of body mass, or loads on their distal legs equal to a total of 5% of body mass. We estimated muscle energy use by measuring blood flow to all of the leg muscles using injected microspheres. Total blood flow to the leg muscles increased by approximately 15% under both loading conditions, which matched the percentage increase in net organismal metabolic rate. Significant increases in energy use(inferred from blood flow) above that found in unloaded birds were found in 12 muscles in trunk-loaded birds, with most of the increases restricted to stance-phase muscles, as predicted. Just three of these muscles, the femerotibialis, the iliotibialis lateralis pars postacetabularis and the fibularis longus accounted for 70% of the increased energy use. Noticeably absent from the group of muscles that increased energy use during trunk loading were several large biarticular muscles that have extensor actions at the hip or ankle, but flexor actions at the knee. We concluded that the low energetic cost of carrying trunk loads in guinea fowl may rely on the activation of a group of muscles that together provide support and propulsion across all the major joints, without producing opposing moments at other joints that could potentially waste energy. The specific leg muscles responsible for the increase in metabolism during trunk loading also suggest that the energy cost of producing mechanical work may be an important determinant of the cost of carrying extra mass on the trunk. During distal-limb loading, eleven leg muscles had significant increases in energy use, but unlike during trunk loading, both stance- and swing-phase muscles had large increases in energy use. This distribution of energy use between stance and swing agrees with the prediction that increased mechanical work determines the cost of limb loading, because a substantial fraction of the increased segmental work during distal-limb loading in guinea fowl has been found to occur during stance.

SUMMARYWe examined the energetic cost of loading the trunk or distal portion of the leg in walking and running guinea fowl (Numida meleagris). These different loading regimes were designed to separately influence the energy use by muscles used during the stance and swing phases of the stride. Metabolic rate, estimated from oxygen consumption, was measured while birds locomoted on a motorized treadmill at speeds from 0.5 to 2.0 m s-1, either unloaded, or with a mass equivalent to 23% of their body mass carried on their backs, or with masses equal to approximately 2.5% of their body mass attached to each tarsometatarsal segment. In separate experiments, we also measured the duration of stance and swing in unloaded, trunk-loaded, or limb-loaded birds. In the unloaded and limb-loaded birds, we also calculated the mechanical energy of the tarsometatarsal segment throughout the stride.Trunk and limb loads caused similar increases in metabolic rate. During trunk loading, the net metabolic rate (gross metabolic rate - resting metabolic rate) increased by 17% above the unloaded value across all speeds. This percentage increase is less than has been found in most studies of humans and other mammals. The economical load carriage of guinea fowl is consistent with predictions based on the relative cost of the stance and swing phases of the stride in this species. However, the available comparative data and considerations of the factors that determine the cost of carrying extra mass lead us to the conclusion that the cost of load carrying is unlikely to be a reliable indicator of the distribution of energy use in stance and swing. Both loading regimes caused small changes in the swing and/or stance durations, but these changes were less than 10%.Loading the tarsometatarsal segment increased its segmental energy by 4.1 times and the segmental mechanical power averaged over the stride by 3.8 times. The increases in metabolism associated with limb loading appear to be linked to the increases in mechanical power. The delta efficiency (change in mechanical power divided by the change in metabolic power) of producing this power increased from 11% in walking to approximately 25% in running. Although tarsometatarsal loading was designed to increase the mechanical energy during swing phase, 40% of the increase in segmental energy occurred during late stance. Thus, the increased energy demand of distal limb loading in guinea fowl is predicted to cause increases in energy use by both stance- and swing-phase muscles.

Estimation of muscle parameters specifying force-length and force-velocity behavior requires in general a large number of sophisticated experiments often including a combination of isometric, isokinetic, isotonic, and quick-release experiments. This study validates a simpler method (ISOFIT) to determine muscle properties by fitting a Hill-type muscle model to a set of isovelocity data. Muscle properties resulting from the ISOFIT method agreed well with muscle properties determined separately in in vitro measurements using frog semitendinosus muscles. The force-length curve was described well by the results of the model. The force-velocity curve resulting from the model coincided with the experimentally determined curve above approximately 20% of maximum isometric force (correlation coefficient R>0.99). At lower forces and thus higher velocities the predicted curve underestimated velocity. The stiffness of the series elastic component determined with direct experiments was approximately 10% lower than that determined by the ISOFIT method. Use of the ISOFIT method can decrease experimental time up to 80% and reduce potential changes in muscle parameters due to fatigue.

Running and walking are mechanically complex activities. Leg muscles must exert forces to support weight and provide stability, do work to accelerate the limbs and body centre of mass, and absorb work to act as brakes. Current understanding of energy use during legged locomotion has been limited by the lack of measurements of energy use by individual muscles. Our study is based on the correlation between blood flow and aerobic energy expenditure in active skeletal muscle during locomotion. This correlation is strongly supported by the available evidence concerning control of blood flow to active muscle, and the relationship between blood flow and the rate of muscle oxygen consumption. We used injectable microspheres to measure the blood flow to the hind-limb muscles, and other body tissues, in guinea fowl (Numida meleagris) at rest, and across a range of walking and running speeds. Combined with data concerning the various mechanical functions of the leg muscles, this approach has enabled the first direct estimates of the energetic costs of some of these functions. Cardiac output increased from 350 ml min(-1) at rest, to 1700 ml min(-1) at a running speed ( approximately 2.6 m s(-1)) eliciting a of 90% of . The increase in cardiac output was achieved via approximately equal factorial increases in heart rate and stroke volume. Approximately 90% of the increased cardiac output was directed to the active muscles of the hind limbs, without redistribution of blood flow from the viscera. Values of mass-specific blood flow to the ventricles, approximately 15 ml min(-1) g(-1), and one of the hind-limb muscles, approximately 9 ml min(-1) g(-1), were the highest yet recorded for blood flow to active muscle. The patterns of increasing blood flow with increasing speed varied greatly among different muscles. The increases in flow correlated with the likely fibre type distribution of the muscles. Muscles expected to have many high-oxidative fibres preferentially increased flow at low exercise intensities. We estimated substantial energetic costs associated with swinging the limbs, co-contraction to stabilize the knee and work production by the hind-limb muscles. Our data provide a basis for evaluating hypotheses relating the mechanics and energetics of legged locomotion.

Explaining the energetics of walking and running has been difficult because the distribution of energy use among individual muscles has not been known. We estimated energy use by measuring blood flow to the hindlimb muscles in guinea fowl. Blood flow to skeletal muscles is controlled locally and varies directly with metabolic rate. We estimate that the swing-phase muscles consume 26% of the energy used by the limbs and the stance-phase muscles consume the remaining 74%, independent of speed. Thus, contrary to some previous suggestions, swinging the limbs requires an appreciable fraction of the energy used during terrestrial legged locomotion. Models integrating the energetics and mechanics of running will benefit from more detailed information on the distribution of energy use by the muscles.

Eels swim in the anguilliform mode in which the majority of the body axis undulates to generate thrust. For this reason, muscle function has been hypothesised to be relatively uniform along the body axis relative to some other teleosts in which the caudal fin is the main site of thrust production. The European eel (Anguilla anguilla L.) has a complex life cycle involving a lengthy spawning migration. Prior to migration, there is a metamorphosis from a yellow (non-migratory) to a silver (migratory) life-history phase. The work loop technique was used to determine slow muscle power outputs in yellow- and silver-phase eels. Differences in muscle properties and power outputs were apparent between yellow- and silver-phase eels. The mass-specific power output of silver-phase slow muscle was greater than that of yellow-phase slow muscle. Maximum slow muscle power outputs under approximated in vivo conditions were 0.24 W kg(−)(1) in yellow-phase eel and 0.74 W kg(−)(1) in silver-phase eel. Power output peaked at cycle frequencies of 0.3-0.5 Hz in yellow-phase slow muscle and at 0.5-0.8 Hz in silver-phase slow muscle. The time from stimulus offset to 90 % relaxation was significantly greater in yellow- than in silver-phase eels. The time from stimulus onset to peak force was not significantly different between life-history stages or axial locations. Yellow-phase eels shifted to intermittent bursts of higher-frequency tailbeats at a lower swimming speed than silver-phase eels. This may indicate recruitment of fast muscle at low speeds in yellow-phase eels to compensate for a relatively lower slow muscle power output and operating frequency.

The Pacific bonito, Sarda chiliensis, is anatomically intermediate between mackerel and tuna. The specialisations exhibited by tuna are present in the bonito, but to a lesser degree. Slow-twitch muscle strain and activity patterns were determined during steady swimming (tailbeat frequency 1.2-3.2 Hz) at four locations on the body of Sarda chiliensis using sonomicrometry and electromyography. Both strain and the phase of electromygraphic activity were independent of tailbeat frequency. The strain of superficial slow-twitch muscle increased from +/−3.1 % l(0) at 0.35FL to +/−5.8 % l(0) at 0.65FL, where l(0) is muscle resting length and FL is the body length from snout to tail fork. Between 0.35 and 0.65FL, there was a negative phase shift of 16 degrees in the onset of electromygraphic activity in superficial slow-twitch muscle relative to the strain cycle. Muscle activity patterns are comparable with those of tuna. At 0.58FL, the onset of activity in deep slow-twitch muscle was approximately synchronous with the onset of activity in superficial muscle in the same myotome at 0.65FL. The distribution of slow-twitch muscle along the body of Sarda chiliensis and four additional fish species, Anguilla anguilla, Oncorhynchus mykiss, Scomber scombrus and Thunnus albacares, was also measured. Slow-twitch muscle appears to become more concentrated at approximately 0.5FL as swimming kinematics become more thunniform.

Undulatory swimming in fish is powered by the segmental body musculature of the myotomes. Power generated by this muscle and the interactions between the fish and the water generate a backward-travelling wave of lateral displacement of the body and caudal fin. The body and tail push against the water, generating forward thrust. The muscle activation and strain patterns that underlie body bending and thrust generation have been described for a number of species and show considerable variation. This suggests that muscle function may also vary among species. This variation must be due in large part to the complex interactions between muscle mechanical properties, fish body form, swimming mode, swimming speed and phylogenetic relationships. Recent work in several laboratories has been directed at studying patterns of muscle power output in vitro under simulated swimming conditions. This work suggests that the way that fish generate muscle power and convert it into thrust through the body and caudal fin does indeed vary. However, despite the differences, several features appear to be common to virtually all species studied and suggest where future effort should be directed if muscle function in swimming fish is to be better understood.

but they are ineYcient at transmitting water, both because they have narrow lumens and because water can only The resistances of xylem vessel walls and perforation pass between them through the primary cell wall. Xylem plates has been investigated using 18 large-scale vessel elements, meanwhile, have evolved from tracheids physical models made of plastic tubing into which through widening of the lumen, shortening of the cell, scale models of plates were inserted. Flow of water and partial loss of the end walls ( Frost, 1930a). Together, through vessels was modelled using glycerol instead these changes have reduced the flow resistance of xylem of water to keep the Reynolds number below 0.1. The vessels, allowing them to conduct water more eYciently technique proved easy, cheap and reliable. up the plant. Results showed that perforation plate resistance is Wider cells are more eYcient because the resistance of low compared with the resistance of the walls, what- a cylindrical vessel is inversely proportional to the fourth ever the plate morphology; plates only provided power of its radius. Resistance, R, is given by the Hagen0.6‐18.6% extra resistance. Simple plates provided Poiseuille equation ( Vogel, 1994) less resistance than scalariform plates, but because they are arranged closer together in vessels, resistance values (1.7‐5.1%) overlap with those of scalari

Bluegill Lepomis macrochirus showed variation in their diet and trophic morphology based on habitat. Pelagic L. macrochirus feed almost exclusively on cladocerans; littoral L. macrochirus feed on a variety of benthic invertebrates, molluscs, cladocerans and insects. Fish from the littoral habitat had wider pharyngeal jaws, which probably allowed them to crush gastropods and process benthic invertebrates.

Swimming flumes enable fish swimming behavior, physiology, and performance to be quantified in ways that are not practicable for fish swimming through open water. By placing fish in a water flow, speed can be controlled, fish can be instrumented to monitor a wide range of physiological parameters, and the exchange of materials between the fish and water can be quantified. This can provide vital information regarding fish fitness and health. If meaningful data are to be obtained, however, careful consideration must be given to flume design and operation, experimental protocol and the physiological state of the fish. Modifications to standard flume designs can potentially allow for accommodation of a wider range of species and experimental conditions that will enhance basic understanding of fish physiology and behavior and can potentially be applied in optimizing aquacultural techniques.

The obliquely striated muscle in the leech body wall has a broad functional repertoire; it provides power for both locomotion and suction feeding. It also operates over an unusually high strain range, undergoing up to threefold changes in length. Serotonin (5-HT) may support this functional flexibility, integrating behavior and biomechanics. It can act centrally, promoting motor outputs that drive body wall movements, and peripherally, modulating the mechanical properties of body wall muscle. During isometric contractions 5-HT enhances active force production and reduces resting muscle tone. We therefore hypothesized that 5-HT would increase net work output during the cyclical contractions associated with locomotion and feeding. Longitudinal strains measured during swimming, crawling and feeding were applied to body wall muscle in vitro with the timing and duration of stimulation selected to maximize net work output. The net work output during all simulated behaviors significantly increased in the presence of 100 μM 5-HT relative to the 5-HT-free control condition. Without 5-HT the muscle strips could not achieve a net positive work output during simulated swimming. The decrease in passive tension associated with 5-HT may also be important in reducing muscle antagonist work during longitudinal muscle lengthening. The behavioral and mechanical effects of 5-HT during locomotion are clearly complementary, promoting particular behaviors and enhancing muscle performance during those behaviors. Although 5-HT can enhance muscle mechanical performance during simulated feeding, low in vivo activity in serotonergic neurons during feeding may mean that its mechanical role during this behavior is less important than during locomotion.

The iliotibialis lateralis pars postacetabularis (ILPO) is the largest muscle in the hindlimb of the guinea fowl and is thought to play an important role during the stance phase of running, both absorbing and producing work. Using sonomicrometry and electromyography, we examined whether the ILPO experiences differential strain between proximal, central and distal portions of the posterior fascicles. When the ILPO is being lengthened while active, the distal portion was found to lengthen significantly more than either the proximal or central portions of the muscle. Our data support the hypothesis that the distal segment lengthened farther and faster because it began activity at shorter sarcomere lengths on the ascending limb of the length–tension curve. Probably because of the self-stabilizing effects of operating on the ascending limb of the length–tension curve, all segments reached the end of lengthening and started shortening at the same sarcomere length. During shortening, this similarity in sarcomere length among the segments was maintained, as predicted from force–velocity effects, and shortening strain was similar in all segments. The differential active strain during active lengthening is thus ultimately determined by differences in strain during the passive portion of the cycle. The sarcomere lengths of all segments of the fascicles were similar at the end of active shortening, but after the passive portion of the cycle the distal segment was shorter. Differential strain in the segments during the passive portion of the cycle may be caused by differential joint excursions at the knee and hip acting on the ends of the muscle and being transmitted differentially by the passive visco-elastic properties of the muscle. Alternatively, the differential passive strain could be due to the action of active or passive muscles in the thigh that transmit force to the IPLO in shear. Based on basic sarcomere dynamics we predict that differential strain is more likely to occur in muscles undergoing active lengthening at the beginning of contraction than those undergoing only shortening. * ILPO : iliotibialis lateralis pars postacetabularis L : the length at the middle of the plateau region of the length–tension curve (optimal length) L max,st : maximum length of the ILPO occurring during stance L max,sw : maximum length of the ILPO occurring during swing L min,fd : minimum length of the ILPO that occurred near the time of foot-down L min,to : minimum length of the ILPO that occurred near the time of toe-off L off : length of the ILPO 25 ms after the stop of the EMG activity L on : length of the ILPO 25 ms after the start of the EMG activity L sc : in vivo sarcomere lengths L sc,ref : reference sarcomere length L so : in vivo segment lengths measured by sonomicrometry L so,ref : sonomicrometer length t al : duration of active lengthening V al : average velocity during active lengthening V as : velocity during active shortening ΔL : passive shortening Δ L al : active lengthening Δ L as : active shortening Δ L as : active shortening Δ L pl : passive lengthening

SUMMARYPhysiological and anatomical evidence suggests that in birds the iliotibialis lateralis pars postacetabularis (ILPO) is functionally important for running. Incorporating regional information, we estimated the mean sarcomere strain trajectory and electromyographic (EMG) amplitude of the ILPO during level and incline walking and running. Using these data and data in the literature of muscle energy use, we examined three hypotheses: (1) active lengthening will occur on the ascending limb of the length–tension curve to avoid potential damage caused by stretch on the descending limb; (2) the active strain cycle will shift to favor active shortening when the birds run uphill and shortening will occur on the plateau and shallow ascending limb of the length–tension curve; and (3) measures of EMG intensity will correlate with energy use when the mechanical function of the muscle is similar. Supporting the first hypothesis, we found that the mean sarcomere lengths at the end of active lengthening during level locomotion were smaller than the predicted length at the start of the plateau of the length–tension curve. Supporting the second hypothesis, the magnitude of active lengthening decreased with increasing slope, whereas active shortening increased. In evaluating the relationship between EMG amplitude and energy use (hypothesis 3), we found that although increases in EMG intensity with speed, slope and loading were positively correlated with muscle energy use, the quantitative relationships between these variables differed greatly under different conditions. The relative changes in EMG intensity and energy use by the muscle probably varied because of changes in the mechanical function of the muscle that altered the ratio of muscle energy use to active muscle volume. Considering the overall function of the cycle of active lengthening and shortening of the fascicles of the ILPO, we conclude that the function of active lengthening is unlikely to be energy conservation and may instead be related to promoting stability at the knee. The work required to lengthen the ILPO during stance is provided by co-contracting knee flexors. We suggest that this potentially energetically expensive co-contraction serves to stabilize the knee in early stance by increasing the mechanical impedance of the joint.

UNLABELLED PREMISE OF THE STUDY Seed dispersal performance is an essential component of plant fitness. Despite their significance in shaping performance, the mechanical processes that drive dispersal are poorly understood. We have quantified seed dispersal mechanics in Cardamine parviflora (Brassicaceae), a ballistic disperser that launches seeds with specialized catapult-like structures. To determine which aspects of catapult function dictate interspecific dispersal differences, we compared this disperser with other ballistic dispersers. Comparison with brassicas that lack ballistic dispersal may also provide insight into the evolution of this mechanism. • METHODS Catapult performance was quantified using high-speed video analysis of dehiscence, ballistic modeling of seed trajectories, and measuring the mechanical energy storage capacity of the spring-like siliqua valve tissue that launched the seeds. • KEY RESULTS The siliquae valves coiled rapidly outward, launching the seeds in 4.7 ± 1.3 ms (mean ± SD, N = 11). Coiling was likely driven by the bilayered valve structure. The catapult was 21.3 ± 10.3% efficient (mean ± SD, N = 11) at transferring stored elastic energy to the seeds as kinetic energy. The majority of seeds (71.4%) were not launched effectively. • CONCLUSIONS The efficiency of the C. parviflora catapult was high in comparison to that of a ballistic diplochore, a dispersal mode associated with poor ballistic performance, although the unreliability of the launch mechanism limited dispersal distance. Effective launching requires temporary seed-valve adhesion. The adhesion mechanism may be the source of the unreliability. Valve curvature is likely driven by the bilayered valve structure, a feature absent in nonballistic brassicas.

Food processing is costly, potentially limiting the energy and time devoted to other essential functions such as locomotion or reproduction. In ectotherms, post-prandial thermophily, the selection of a warm environmental temperature after feeding, may be advantageous in minimizing the duration of this elevated cost. Although present in many vertebrate taxa, this behaviour had not previously been observed in invertebrates. Sanguivorous leeches ingest large blood meals that are costly to process and limit mobility until excess fluid can actively be expelled to reduce body volume. When presented with a temperature gradient from 10°C to 30°C, leeches select a temperature that is significantly warmer (24.3 ± 0.9°C,n= 6) than their acclimation temperature (Ta, 21°C). Unfed leeches preferred temperatures that were significantly cooler than ambient (12.8 ± 0.9°C,n= 6). This behavioural strategy is consistent with minimizing the time course of elevated post-feeding energy costs and reducing energy expenditure during fasting. Our observations raise the possibility that thermoregulatory behaviour of this type is an unrecognized feature of other invertebrate taxa.

Many fish transfer mechanical power to the surrounding water using lateral waves of body axis undulation. Waves of curvature progress along the axis in a rostrocaudal direction. These are produced by rostrocaudally progressing waves of muscle activity driving sequential contraction of the myotomal muscle blocks. Muscle activity is offset on opposite sides of the body. Regions of active muscle produce local shortening while active, contralateral muscle on the outside of the bend is stretched while inactive. Muscle force is transferred posteriorly to the axial skeleton and caudal fin by connective tissue and skeletal elements.

[Figure][1] David Ellerby discusses Paul Webb's 1971 paper entitled ‘The swimming energetics of Trout. II. Oxygen consumption and swimming efficiency’. A copy of the paper can be obtained from Compared with other modes of

The effects of acute temperature change on the cost of bluegill Lepomis macrochirus swimming were quantified. At 14 degrees C, maximum labriform swimming speed (U(lab,max)) was reduced relative to that at the acclimation temperature of 22 degrees C, but total cost of transport (T(TC)) remained unchanged. At 30 degrees C, U(lab,max) was the same as at 22 degrees C, but T(TC) was 66% greater.

Explosive dehiscence ballistically disperses seeds in a number of plant species. During dehiscence, mechanical energy stored in specialized tissues is transferred to the seeds to increase their kinetic and potential energies. The resulting seed dispersal patterns have been investigated in some ballistic dispersers, but the mechanical performance of a launch mechanism of this type has not been measured. The properties of the energy storage tissue and the energy transfer efficiency of the launch mechanism were quantified in Impatiens capensis. In this species the valves forming the seed pod wall store mechanical energy. Their mass specific energy storage capacity (124 J kg(-1)) was comparable with that of elastin and spring steel. The energy storage capacity of the pod tissues was determined by their level of hydration, suggesting a role for turgor pressure in the energy storage mechanism. During dehiscence the valves coiled inwards, collapsing the pod and ejecting the seeds. Dehiscence took 4.2+/-0.4 ms (mean +/-SEM, n=13). The estimated efficiency with which energy was transferred to the seeds was low (0.51+/-0.26%, mean +/-SEM, n=13). The mean seed launch angle (17.4+/-5.2, mean +/-SEM, n=45) fell within the range predicted by a ballistic model to maximize dispersal distance. Low ballistic dispersal efficiency or effectiveness may be characteristic of species that also utilize secondary seed dispersal mechanisms.

SUMMARYThe pectoralis muscles are the main source of mechanical power for avian flight. The power output of these muscles must be modulated to meet the changing power requirements of flight across a range of speeds. This can be achieved at the muscle level by manipulation of strain trajectory and recruitment patterns, and/or by intermittent flight strategies. We have measured the in vitro power outputs of pectoralis muscle fascicles from budgerigars Melopsittacus undulatus and zebra finches Taeniopygia guttata under conditions replicating those previously measured in vivo during flight. This has allowed us to quantify the extent to which different power modulation mechanisms control flight muscle power output. Intermittent flight behaviour is a more important determinant of flight power in zebra finches than budgerigars. This behaviour accounts for 25–62% of power modulation relative to the maximum available mechanical power output in zebra finch, compared to 0–38% in budgerigars. Muscle level changes in fascicle strain trajectory and motor unit recruitment, rather than intermittent flight behaviours, are the main determinants of pectoralis muscle power output in budgerigars at all speeds, and in zebra finch at speeds below 14 m s–1.

SUMMARYFlight power varies in a U-shaped relationship with flight speed, requiring the modulation of flight muscle power in order to meet these changing power demands. The power output of the pectoralis muscle can potentially be modulated by changing strain trajectory and the relative timing and intensity of muscle activity. Pectoralis muscle length change and activity patterns were recorded in budgerigars Melopsitaccus undulatus and zebra finches Taeniopygia guttata at a range of flight speeds using sonomicrometry and electromyography (EMG). The pectoralis muscles in these species contain a single muscle fibre type. Therefore, the power output is entirely determined by muscle activity and strain trajectory, rather than recruitment of motor units with different contractile properties as in many other vertebrate muscle systems. Relative EMG intensity, wingbeat frequency and muscle strain varied in an approximately U-shaped relationship with flight speed. The shape of the length trajectory varied with flight speed in budgerigars, with the proportion of the cycle spent shortening being lowest at intermediate flight speeds. In zebra finch pectoralis muscle the shape of the length trajectory did not vary significantly with flight speed. In both species the observed changes in muscle recruitment and length trajectory are consistent with meeting flight power requirements that vary in a U-shaped pattern with speed. Both species utilised intermittent flight, tending to spend relatively less time flapping at intermediate flight speeds. This supports the idea that intermittent flight is used as a simple power modulation strategy. However, the idea that intermittent flight serves to maintain a `fixed gear' is over-simplistic and fails to recognise the plasticity in performance at the level of the muscle. Intermittent flight is only one component of a complex power modulation strategy.

SUMMARYBluegill sunfish (Lepomis macrochirus) swim in the labriform mode at low speeds, generating lift and thrust by beating their pectoral fins. The maximal power output available from the two largest pectoral fin adductor and abductor muscles, constituting half of the total pectoral girdle muscle mass,was measured in vitro and used to estimate the muscle mechanical power output during maximal labriform swimming (Pmech;0.15–0.21 W kg–1 body mass). Respirometry was used to estimate the total metabolic power input (Ptotal; 0.95 W kg–1 body mass) and the metabolic power available to the active muscle mass (Pmuscle; Ptotalminus standard metabolic rate, 0.57 W kg–1 body mass) at this swimming speed. Drag measurements made on towed, dead fish were used to estimate the mechanical power required to overcome body drag(Pdrag; 0.028 W kg–1 body mass). Efficiency estimates based on these data fell into the following ranges:overall swimming efficiency(ηgross=Pmech/Ptotal),0.16–0.22; muscle efficiency(ηmuscle=Pmech/Pmuscle),0.26–0.37; and propeller efficiency(ηprop=Pdrag/Pmech),0.15–0.20. Comparison with other studies suggests that labriform swimming may be more efficient than swimming powered by undulations of the body axis.

A major goal of flight research has been to establish the relationship between the mechanical power requirements of flight and flight speed. This relationship is central to our understanding of the ecology and evolution of bird flight behaviour. Current approaches to determining flight power have relied on a variety of indirect measurements and led to a controversy over the shape of the power–speed relationship and a lack of quantitative agreement between the different techniques. We have used a new approach to determine flight power at a range of speeds based on the performance of the pectoralis muscles. As such, our measurements provide a unique dataset for comparison with other methods. Here we show that in budgerigars (Melopsittacus undulatus) and zebra finches (Taenopygia guttata) power is modulated with flight speed, resulting in U-shaped power–speed relationship. Our measured muscle powers agreed well with a range of powers predicted using an aerodynamic model. Assessing the accuracy of mechanical power calculated using such models is essential as they are the basis for determining flight efficiency when compared to measurements of flight metabolic rate and for predicting minimum power and maximum range speeds, key determinants of optimal flight behaviour in the field.

We examined the energetic cost of loading the trunk or distal portion of the leg in walking and running guinea fowl (Numida meleagris). These different loading regimes were designed to separately influence the energy use by muscles used during the stance and swing phases of the stride. Metabolic rate, estimated from oxygen consumption, was measured while birds locomoted on a motorized treadmill at speeds from 0.5 to 2.0 m s-1, either unloaded, or with a mass equivalent to 23% of their body mass carried on their backs, or with masses equal to approximately 2.5% of their body mass attached to each tarsometatarsal segment. In separate experiments, we also measured the duration of stance and swing in unloaded, trunk-loaded, or limb-loaded birds. In the unloaded and limb-loaded birds, we also calculated the mechanical energy of the tarsometatarsal segment throughout the stride. Trunk and limb loads caused similar increases in metabolic rate. During trunk loading, the net metabolic rate (gross metabolic rate-resting metabolic rate) increased by 17% above the unloaded value across all speeds. This percentage increase is less than has been found in most studies of humans and other mammals. The economical load carriage of guinea fowl is consistent with predictions based on the relative cost of the stance and swing phases of the stride in this species. However, the available comparative data and considerations of the factors that determine the cost of carrying extra mass lead us to the conclusion that the cost of load carrying is unlikely to be a reliable indicator of the distribution of energy use in stance and swing. Both loading regimes caused small changes in the swing and/or stance durations, but these changes were less than 10%. Loading the tarsometatarsal segment increased its segmental energy by 4.1 times and the segmental mechanical power averaged over the stride by 3.8 times. The increases in metabolism associated with limb loading appear to be linked to the increases in mechanical power. The delta efficiency (change in mechanical power divided by the change in metabolic power) of producing this power increased from 11% in walking to approximately 25% in running. Although tarsometatarsal loading was designed to increase the mechanical energy during swing phase, 40% of the increase in segmental energy occurred during late stance. Thus, the increased energy demand of distal limb loading in guinea fowl is predicted to cause increases in energy use by both stance- and swing-phase muscles.