Intra-articular anaesthesia mitigates established pain in experimental osteoarthritis: a preliminary study of gait impulse redistribution as a biomarker of analgesia pharmacodynamics.

OBJECTIVE: Develop a sensitive, functional biomarker of persistent joint pain in a large animal model of experimental osteoarthritis. Evaluate Impulse Ratio as a measure of weight distribution among supporting limbs throughout the early natural history of osteoarthritis and with local anaesthesia and analgesia. DESIGN: The distribution of weight bearing in the trot of 11 skeletally-mature dogs was analyzed before and after unilateral surgical intervention (cranial cruciate transection or distal femoral focal impact). The short-term effects of two analgesic treatments (intra-articular lidocaine and intra-dermal meloxicam) were then evaluated as an index of pain relief based on the redistribution of weight-bearing impulse between normal and injured limbs. RESULTS: Impulse Ratio was able to resolve weight redistribution between limbs in both long-term (weekly for over 400 days) and short-term (15 min intervals) joint evaluations. Joint pain relief from lidocaine administration could be reliably tracked over its brief acting time course. Meloxicam administration resulted in ambiguous results, where average weight bearing in the injured limb did not increase, but the variability of limb use changed transiently and reversibly. CONCLUSION: Joint function and the role of persistent joint pain in the development of osteoarthritis can be investigated effectively and efficiently in a large animal model through the use of Impulse Ratio. Impulse Ratio can be a functionally relevant and sensitive biomarker of locomotion-related joint pain.

Contractile properties of muscle fibers from the deep and superficial digital flexors of horses.

Equine digital flexor muscles have independent tendons but a nearly identical mechanical relationship to the main joint they act upon. Yet these muscles have remarkable diversity in architecture, ranging from long, unipennate fibers ("short" compartment of DDF) to very short, multipennate fibers (SDF). To investigate the functional relevance of the form of the digital flexor muscles, fiber contractile properties were analyzed in the context of architecture differences and in vivo function during locomotion. Myosin heavy chain (MHC) isoform fiber type was studied, and in vitro motility assays were used to measure actin filament sliding velocity (V(f)). Skinned fiber contractile properties [isometric tension (P(0)/CSA), velocity of unloaded shortening (V(US)), and force-Ca(2+) relationships] at both 10 and 30°C were characterized. Contractile properties were correlated with MHC isoform and their respective V(f). The DDF contained a higher percentage of MHC-2A fibers with myosin (heavy meromyosin) and V(f) that was twofold faster than SDF. At 30°C, P(0)/CSA was higher for DDF (103.5 ± 8.75 mN/mm(2)) than SDF fibers (81.8 ± 7.71 mN/mm(2)). Similarly, V(US) (pCa 5, 30°C) was faster for DDF (2.43 ± 0.53 FL/s) than SDF fibers (1.20 ± 0.22 FL/s). Active isometric tension increased with increasing Ca(2+) concentration, with maximal Ca(2+) activation at pCa 5 at each temperature in fibers from each muscle. In general, the collective properties of DDF and SDF were consistent with fiber MHC isoform composition, muscle architecture, and the respective functional roles of the two muscles in locomotion.

Contractile behavior of the forelimb digital flexors during steady-state locomotion in horses (Equus caballus): an initial test of muscle architectural hypotheses about in vivo function.

The forelimb digital flexors of the horse display remarkable diversity in muscle architecture despite each muscle-tendon unit having a similar mechanical advantage across the fetlock joint. We focus on two distinct muscles of the digital flexor system: short compartment deep digital flexor (DDF(sc)) and the superficial digital flexor (SDF). The objectives were to investigate force-length behavior and work performance of these two muscles in vivo during locomotion, and to determine how muscle architecture contributes to in vivo function in this system. We directly recorded muscle force (via tendon strain gauges) and muscle fascicle length (via sonomicrometry crystals) as horses walked (1.7 m s(-1)), trotted (4.1 m s(-1)) and cantered (7.0 m s(-1)) on a motorized treadmill. Over the range of gaits and speeds, DDF(sc) fascicles shortened while producing relatively low force, generating modest positive net work. In contrast, SDF fascicles initially shortened, then lengthened while producing high force, resulting in substantial negative net work. These findings suggest the long fibered, unipennate DDF(sc) supplements mechanical work during running, whereas the short fibered, multipennate SDF is specialized for economical high force and enhanced elastic energy storage. Apparent in vivo functions match well with the distinct architectural features of each muscle.

Superficial digital flexor tendon lesions in racehorses as a sequela to muscle fatigue: a preliminary study.

REASON FOR PERFORMING STUDY: Racing and training related lesions of the forelimb superficial digital flexor tendon are a common career ending injury to racehorses but aetiology and/or predisposing causes of the injury are not completely understood. OBJECTIVES: Although the injury takes place within the tendon, the lesion must be considered within the context of the function of the complete suspensory system of the distal limb, including the associated muscles. METHODS: Both muscle and tendon function were investigated in vivo using implanted strain gauges in 3 Thoroughbred horses walking, trotting and cantering on a motorised treadmill. These data were combined with assessments of muscle architecture and fibre composition to arrive at an overview of the contribution of each muscle-tendon unit during locomotion. RESULTS: The superficial digital flexor muscle has fatigue-resistant and high force production properties that allow its tendon to store and return elastic energy, predominantly at the trot. As running speed increases, deep digital flexor tendon force increases and it stabilises hyperextension of the fetlock, thus reinforcing the superficial digital flexor in limb load support. The deep digital flexor muscle has fast contracting properties that render it susceptible to fatigue. CONCLUSION: Based on these measurements and supporting evidence from the literature, it is proposed that overloading of the superficial digital flexor tendon results from fatigue of the synergistic, faster contracting deep digital flexor muscle. POTENTIAL RELEVANCE: Future research investigating distal limb system function as a whole should help refine clinical diagnostic procedures and exercise training approaches that will lead to more effective prevention and treatment of digital flexor tendon injuries in equine athletes.

A novel resource model for underprivileged health support: Community Medical Outreach.

Community Medical Outreach is a student-run organization that provides healthcare access to medically underprivileged farm workers. The program exploits the substantial energy, enthusiasm and organizational capacity of pre-medical students as they prepare to apply for medical school. All of the partners benefit from the interchange. The students gain from a unique first-hand medical experience that demonstrates their leadership, management skills, commitment to a healthcare team, and focus on care at the community level. Those in most need gain healthcare access. Volunteer staff and physicians are inspired by the students and are energized by caring for those most in need of health care. The companies, agencies, and organizations donating supplies, drugs, expertise, and sponsorship benefit from enhanced public relations. The article describes the initiation, the lessons learned, the critical importance of linkages, and the essential components such as individual and institutional liability. Community Medical Outreach is an important vehicle for shaping students in the process of becoming physicians, shaping those involved with the process of admitting students, shaping training experiences, and shaping new models of health care.

Morphology, histochemistry, and function of epaxial cervical musculature in the horse (Equus caballus).

The semispinalis capitis and splenius muscles of the horse were analyzed for gross morphology, microarchitecture, fiber length, and fiber type. Although these two muscles are similar in size and anatomical position, they are very different from one another in structural design and histochemistry, implying diverse functional roles in the animal's behavior. The histochemical staining profile was limited to two fiber types: slow oxidative and fast glycolytic. The splenius muscle has simple architecture, long fibers, and a 60/40 ratio of SO to FG cross-sectional area. The semispinalis capitis has complex architecture with short-fibered, concentric compartments dorsal to its central tendon and longer-fibered compartments ventrally. The entire dorsal region has an increasing gradient of slow oxidative fiber percentage from caudal to cranial (58-71% SO). In contrast, the ventral region has a decreasing gradient of slow oxidative fibers from caudal to cranial (48-67% FG). These patterns can be interpreted within the context of the cervical musculature during locomotion and posture to indicate the functional advantages of this organization.

Segmental in vivo vertebral kinematics at the walk, trot and canter: a preliminary study.

Understanding the pathophysiology of equine back problems, for clinical evaluation, treatment or injury prevention, requires understanding of the normal 3-dimensional motion characteristics of the vertebral column. Recent studies have investigated regional vertebral kinematics; however, there are no reported measures of direct in vivo segmental vertebral kinematics in exercising horses. Relative movements between 2 adjacent vertebrae were recorded for 3 horses that were clinically sound and did not have a known history of a back problem. A transducer consisting of 2 fixtures and an array of liquid metal strain gauges (LMSGs) was used to measure 3-dimensional segmental vertebral motion. The transducer was attached directly to Steinmann pins implanted in the dorsal spinous processes of adjacent vertebrae in 3 vertebral regions: thoracic (T14 to T16), lumbar (L1 to L3) and lumbosacral (L6 to S2). Rotational displacements between adjacent vertebrae were calculated from the differential outputs of the LMSG array during walk, trot and canter on a treadmill. Peak magnitudes of dorsoventral flexion, lateral bending and axial rotation were recorded continuously for each stride. The largest motion of the 3 instrumented vertebral segments was at the lumbosacral junction. In general, the greatest magnitude of segmental vertebral motion occurred during the canter and the least during the trot. The dynamic and continuous measure of 3-dimensional in vivo segmental vertebral motion provides an important new perspective for evaluating vertebral motion and back problems in horses.

Mechanical energy oscillations of two brachiation gaits: measurement and simulation.

How do arm-swinging apes locomote effectively over a variety of speeds? One way to reduce the metabolic energy cost of locomotion is to transfer energy between reversible mechanical modes. In terrestrial animals, at least two transfer mechanisms have been identified: 1) a pendulum-like mechanism for walking, with exchange between gravitational potential energy and translational kinetic energy, and 2) a spring-like mechanism for running, where the elastic strain energy of stretched muscle and tendon is largely returned to reaccelerate the animal. At slower speeds, a brachiator will always have at least one limb in contact with the support, similar to the overlap of foot contact in bipedal walking. At faster speeds, brachiators exhibit an aerial phase, similar to that seen in bipedal running. Are there two distinct brachiation gaits even though the animal appears to simply swing beneath its overhead support? If so, are different exchange mechanisms employed? Our kinetic analysis of brachiation in a white-handed gibbon (Hylobates lar) indicates that brachiation is indeed comprised of two mechanically distinct gaits. At slower speeds in "continuous contact" brachiation, the gibbon utilizes a simple pendulum-like transfer of mechanical energy within each stride. At faster speeds in "ricochetal" brachiation, translational and rotational kinetic energy are exchanged in a novel "whip-like" transfer. We propose that brachiators utilize the transfer between translational and rotational kinetic energy to control the dynamics of their swing. This maneuver may allow muscle action at the shoulder to control the transfer and adjust the ballistic portion of the step to meet the requirements for the next hand contact.

Multiple walking speed-frequency relations are predicted by constrained optimization.

A person constrained to walk at a given speed v on a treadmill, chooses a particular step frequency f and step length d=v/f. Testing over a range of speeds generates a speed-frequency (v-f) relationship. This relationship is commonly posited as a basic feature of human gait. It is often further posited that this curve follows from minimum energy cost strategy. We observed that individuals walking under different constraint circumstances--walking to a range of fixed metronome frequencies (fixed f) or over a range of spaced markers (fixed d)--produce speed-frequency relations distinct from the constrained v relation. We show here that three distinct speed-frequency curves, similar to those observed, are predicted by the assumption that a walking person optimizes an underlying objective function F (v, f) that has a minimum at the preferred gait. Further, the metabolic cost of transport is a reasonable approximate candidate for the function F.

Fourier analysis of acetabular shape in native American Arikara populations before and after acquisition of horses.

The goal of this study was to identify changes in acetabular morphology associated with the use of horses by Native Americans. Previous studies reported "elongate" acetabula in horseback-riding members of the Omaha and Ponca populations. Such a difference in acetabular shape is a potentially useful osteological marker of habitual horseback riding. This report compares acetabula of adult males from two Native American Arikara populations known to have differed substantially in their use of horses. Population samples were from separate sites in South Dakota: Larson (nonriding) and Leavenworth (riding). Outlines of acetabular rims were digitized and analyzed, using a simplified 12-point Fourier analysis. A Fourier series with six terms accurately described acetabular shape. Significant differences (P<0.10) between riding and nonriding populations were observed in two Fourier coefficients. Acetabula of riding Arikara were found to have smaller B(4) coefficients (P = 0. 061) and more positive B(2) coefficients (P = 0.080), indicating expanded anterior-superior borders relative to acetabula of non-riding Arikara.

External forces and torques generated by the brachiating white-handed gibbon (Hylobates lar).

We compared the kinetics of brachiation to bipedal walking and running. Gibbons use pectoral limbs in continuous contact with their overhead support at slow speeds, but exhibit aerial phases (or ricochetal brachiation) at faster speeds. This basic interaction between limb and support suggests some analogy to walking and running. We quantified the forces in three axes and torque about the vertical axis generated by a brachiating White-handed gibbon (Hylobates lar) and compared them with bipedal locomotion. Handholds oriented perpendicular to the direction of travel (as in ladder rungs) were spaced 0.80, 1.20, 1.60, 1.72, 1.95, and 2.25 m apart. The gibbon proportionally matched forward velocity to stride length. Handhold reaction forces resembled ground reaction forces of running humans except that the order of horizontal braking and propulsion were reversed. Peak vertical forces in brachiation increased with speed as in bipedal locomotion. In contrast to bipedalism, however, peak horizontal forces changed little with speed. Gait transition occurred within the same relative velocity range as the walk-run transition in bipeds (Froude number = 0.3-0.6). We oriented handholds parallel to the direction of travel (as in a continuous pole) at 0.80 and 1.60 m spacings. In ricochetal brachiation, the gibbon generated greater torque with handholds oriented perpendicular as opposed to parallel to the direction of travel. Handhold orientation did not affect peak forces. The similarities and differences between brachiation and bipedalism offer insight into the ubiquity of mechanical principles guiding all limbed locomotion and the distinctiveness of brachiation as a unique mode of locomotion.

Comparison of the trotting gaits of Labrador Retrievers and Greyhounds.

OBJECTIVE: To compare the trotting gaits of Labrador Retrievers and Greyhounds to determine whether differences in locomotion are attributable to differences in their manner of moving or to body size and shape differences between these 2 breeds. ANIMALS: 8 healthy 5-month-old Greyhounds and 5 healthy Labrador Retrievers between 6 and 18 months old. PROCEDURE: A series of 4 force platforms was used to record independent ground reaction forces on the forelimbs and hind limbs during trotting. Values of stride parameters were compared between breeds before and after normalization for size differences. Standard values of absolute and normalized stride period and stride length were determined from linear regressions of these parameters on relative (normalized) velocity. Forces were normalized to body weight and compared at the same relative velocity. RESULTS: Greyhounds used fewer, longer strides than the Labrador Retrievers to travel at the same absolute speed. After normalization for body size differences, most measurable differences between breeds were eliminated. Subtle differences that did persist related to proportion of the stride that the forefoot was in contact with the ground, timing of initial hind foot contact relative to initial forefoot contact, and distribution of vertical force between the forelimbs and hind limbs. CONCLUSIONS AND CLINICAL RELEVANCE: Results suggest that apparent differences in the trotting gait between Labrador Retrievers and Greyhounds are mainly attributable to differences in size, and that dogs of these 2 breeds move in a dynamically similar manner at the trot.

The mechanical behaviour of a novel mammalian intervertebral joint.

The mechanics of mammalian intervertebral joints are complicated by the viscoelastic nature of the connective tissues joining vertebrae, and by multiple vertebral articulations and complex morphologies. Further, interspecific variation in these structures can greatly compound their functional variation between species, making comparative mechanical analyses even more difficult. Despite these sources of variation however, mammalian intervertebral joints universally exhibit a creep relaxation behaviour based on the viscoelastic nature of the soft tissue joint. We have evaluated, in 6 degrees of freedom, the mechanical signature of a novel mammalian lumbar intervertebral joint found in the Scutisorex spine, and compared it with a more typical mammalian joint in the Rattus (rat) lumbar spine. Scutisorex, the hero shrew, is an East African species of shrew with what is likely the most highly modified vertebral morphology in the entire history of mammals. Thus we decided to evaluate the mechanical behaviour of the intervertebral joint of this species, comparing it with a more representative mammal species in Rattus. We built a custom, 6 degrees of freedom, intervertebral joint transducer and a combined axial moment and load application system in order to quantify and compare the complex mechanical behaviour of these joints. Our results suggest that the Scutisorex joint is 5 times more resilient to simple axial torsion per body mass unit than Rattus, and that the complex load (combined axial compression and torsion) mechanical signature of Scutisorex is probably novel among all mammalian intervertebral joints. Under significant but physiological axial compression the Scutisorex intervertebral joint demonstrates no creep relaxation behaviour, simulating the mechanical behaviour of a rigid construct rather than a viscoelastic joint. The purpose of this rigid intervertebral joint in the ecology of Scutisorex remains unknown.

Acceleration and balance in trotting dogs.

During quadrupedal trotting, diagonal pairs of limbs are set down in unison and exert forces on the ground simultaneously. Ground-reaction forces on individual limbs of trotting dogs were measured separately using a series of four force platforms. Vertical and fore-aft impulses were determined for each limb from the force/time recordings. When mean fore-aft acceleration of the body was zero in a given trotting step (steady state), the fraction of vertical impulse on the forelimb was equal to the fraction of body weight supported by the forelimbs during standing (approximately 60 %). When dogs accelerated or decelerated during a trotting step, the vertical impulse was redistributed to the hindlimb or forelimb, respectively. This redistribution of the vertical impulse is due to a moment exerted about the pitch axis of the body by fore-aft accelerating and decelerating forces. Vertical forces exerted by the forelimb and hindlimb resist this pitching moment, providing stability during fore-aft acceleration and deceleration.

A point-mass model of gibbon locomotion.

In brachiation, an animal uses alternating bimanual support to move beneath an overhead support. Past brachiation models have been based on the oscillations of a simple pendulum over half of a full cycle of oscillation. These models have been unsatisfying because the natural behavior of gibbons and siamangs appears to be far less restricted than so predicted. Cursorial mammals use an inverted pendulum-like energy exchange in walking, but switch to a spring-based energy exchange in running as velocity increases. Brachiating apes do not possess the anatomical springs characteristic of the limbs of terrestrial runners and do not appear to be using a spring-based gait. How do these animals move so easily within the branches of the forest canopy? Are there fundamental mechanical factors responsible for the transition from a continuous-contact gait where at least one hand is on a hand hold at a time, to a ricochetal gait where the animal vaults between hand holds? We present a simple model of ricochetal locomotion based on a combination of parabolic free flight and simple circular pendulum motion of a single point mass on a massless arm. In this simple brachiation model, energy losses due to inelastic collisions of the animal with the support are avoided, either because the collisions occur at zero velocity (continuous-contact brachiation) or by a smooth matching of the circular and parabolic trajectories at the point of contact (ricochetal brachiation). This model predicts that brachiation is possible over a large range of speeds, handhold spacings and gait frequencies with (theoretically) no mechanical energy cost. We then add the further assumption that a brachiator minimizes either its total energy or, equivalently, its peak arm tension, or a peak tension-related measure of muscle contraction metabolic cost. However, near the optimum the model is still rather unrestrictive. We present some comparisons with gibbon brachiation showing that the simple dynamic model presented has predictive value. However, natural gibbon motion is even smoother than the smoothest motions predicted by this primitive model.

Biomechanical comparison of two plating techniques for fixation of acetabular osteotomies in dogs.

OBJECTIVE: To compare the failure properties of a 5-hole, 2.7-mm curved acetabular plate (AP) to a 5-hole, 3.5-mm reconstruction plate (RP) when applied to acetabular osteotomies. STUDY DESIGN: Cadaver study. ANIMALS OR SAMPLE POPULATION: Pelves of 8 mature, large-breed dogs. METHODS: A 5-hole, 2.7-mm AP and a 5-hole, 3.5-mm RP were contoured and applied to the dorsal acetabulum of each pelvis. A central acetabular fracture was simulated after plate application by a transverse osteotomy with a fine saw. Each acetabulum was loaded in a weight-bearing direction. A load-deformation curve was produced for each construct, and biomechanical properties of the AP and RP were compared with the Student's paired t-test. A P value of < .05 was considered significant. RESULTS: For the AP and RP composite respectively, the mean +/- SD maximum load to failure was 2,721 +/- 632 N and 2,488 +/- 800 N, the stiffness was 4.8 +/- 1.8 N/m and 5.3 +/- 1.9 N/m, and the energy absorbed was 15.1 +/- 5.2 Nm and 16.3 +/- 8.3 Nm. None of these differences was statistically significant. CONCLUSIONS: Both fixation techniques provided comparable strength, stiffness, and energy absorbed under the loading conditions of this study. CLINICAL RELEVANCE: Because of the relative ease of application, the 2.7-mm curved AP may be the practical choice for acetabular fracture repair in large dogs.

Fractal-based image texture analysis of trabecular bone architecture.

Fractal-based image analysis methods are investigated to extract textural features related to the anisotropic structure of trabecular bone from the X-ray images of cubic bone specimens. Three methods are used to quantify image textural features: power spectrum, Minkowski dimension and mean intercept length. The global fractal dimension is used to describe the overall roughness of the image texture. The anisotropic features formed by the trabeculae are characterised by a fabric ellipse, whose orientation and eccentricity reflect the textural anisotropy of the image. Tests of these methods with synthetic images of known fractal dimension show that the Minkowski dimension provides a more accurate and consistent estimation of global fractal dimension. Tests on bone x-ray (eccentricity range 0.25-0.80) images indicate that the Minkowski dimension is more sensitive to the changes in textural orientation. The results suggest that the Minkowski dimension is a better measure for characterising trabecular bone anisotropy in the x-ray images of thick specimens.

Mechanics of avian fibrous periosteum: tensile and adhesion properties during growth.

We report the results of direct mechanical tests of the fibrous periosteum from the tibiotarsi of white leghorn chicks at 4, 6, 8, 9, 10, 11, 12, and 14 weeks of age using a newly developed sample isolation technique. Additionally, this technique allows the determination of the apparent in vivo load on the fibrous periosteum. The periosteum has a highly nonlinear stress-strain relationship at all ages. For loading below the in vivo level, the periosteum is pliant and mean tensile modulus is 3.35 MPa (+/- 1.84 SD, n = 75). For loading above the in vivo level, tensile stiffness is nearly two orders of magnitude greater. In the region of high stiffness, mean modulus is 229.5 MPa (+/- 89.6, n = 72). In vivo, the periosteum is loaded at the transition between these two stiffness regions. We interpret this as indicating that, in vivo, the collagen fibers of the periosteum are aligned, but subject to minimal loading. Stress levels in the periosteum corresponding to in vivo conditions indicate modest loading, and mean apparent in vivo stress levels are 0.92 MPa (+/- 0.37 SD, n = 67). A second technique demonstrated that the adhesion of the periosteum in the diaphyseal region (1-6 weeks of age) is minimal, but is substantial in the metaphyseal region. The metaphyseal adhesion will affect the transmission of load between the physes. These studies suggest that growth of the fibrous periosteum follows the longitudinal growth of the bone, rather than the periosteum having a direct mechanical influence on growth plate activity. Comparison of tensile properties over the course of growth indicates a substantial increase in periosteal stiffness in the early portion of the growth period, which reaches a maximum at approximately 9 weeks posthatching. There is also a marked decline in periosteal stiffness as growth rate declines in the latest stages of growth (14 weeks). This suggests that the basic properties of periosteal collagen may undergo a transition during the course of this tissue's brief functional lifetime; that is, during long bone growth.

Paralysis and long bone growth in the chick: growth shape trajectories of the pelvic limb.

Growth of chick embryonic femora, tibiotarsi and first phalanges of digit three were measured at one day intervals from day 6 through 16 of incubation. Normal controls were compared to embryos paralyzed at 5 days of incubation. Over the 10 day study period, length of the paralyzed femora, length and width of the paralyzed tibiotarsi and differences in length of the phalanges were observed. Growth in length of phalanx one of digit three was most affected by paralysis over this period. Changes in shape of these bones also occurred during growth. Normal long bones undergo changes in shape as differential growth in length and width occurs. Such changes in shape can be considered as the bone's normal growth "trajectory". Paralyzed bones displayed a different growth trajectory than normal bones. As expected, the long bones of paralyzed embryos were shorter than age-matched controls. Contrary to expectations, however, paralyzed long bones were relatively more stout than age-matched controls.

Mechanical work as a determinant of prey-handling behavior in the tokay gecko (Gekko gecko).

In this study an in vitro analysis of the force and mechanical work required to bite prey items of different size and physical character is combined with an in vivo analysis of prey-handling behavior in the tokay gecko (Gekko gecko). The force required to bite and the work of biting increase with prey size, but the rate of increase is prey specific, with crickets (Acheta domestica) requiring substantially more force and work per bite than larvae (Galleria mellonella and Manduca sexta) for all but the smallest prey. Prey-handling behavior is also prey specific. Geckos exert more bites per feeding event on small crickets than on small insect larvae, but the number of bites increases faster with prey mass for larvae than for crickets. Combination of the in vitro mechanical measurements with the in vivo behavior analysis allows the calculation of total mechanical work per feeding event and indicates that total work increases with prey size but that the difference between prey types is far less than predicted from the differences in structural properties of the prey. This occurs because the number of bites and work per bite relationships tend to cancel the differences in the total work necessary to process each prey type. Thus, when considering the effect of prey size, a 13-fold greater rate of increase in bite force and an 18-fold greater rate of increase of work per bit for crickets over larvae was partially compensated for by a threefold increase in the number of bites used on larvae relative to crickets. These results can be interpreted in two ways. The effect of mechanical work in feeding behavior suggests that the energetics of jaw adductor musculature could play a greater role in governing the feeding behavior of this lizard than has previously been expected. Alternatively, the scaling of work in feeding over a range of prey sizes suggests distinct differences in the geometric features of the prey that determine how they are processed.