Friction between human finger flexor tendons and pulleys at high loads

A method was developed to indirectly measure friction between the flexor tendons and pulleys of the middle and ring finger in vivo. An isokinetic movement device to determine maximum force of wrist flexion, interphalangeal joint flexion (rolling in and out) and isolated proximal interphalangeal (PIP) joint flexion was built. Eccentric and concentric maximum force of these three different movements where gliding of the flexor tendon sheath was involved differently (least in wrist flexion) was measured and compared. Fifty-one hands in 26 male subjects were evaluated. The greatest difference between eccentric and concentric maximum force (29.9%) was found in flexion of the PIP joint. Differences in the rolling in and out movement (26.8%) and in wrist flexion (14.5%) were significantly smaller. The force of friction between flexor tendons and pulleys can be determined by the greater difference between eccentric and concentric maximum force provided by the same muscles in overcoming an external force during flexion of the interphalangeal joints and suggests the presence of a non-muscular force, such as friction. It constitutes of 9% of the eccentric flexion force in the PIP joint and therefore questions the low friction hypothesis at high loads.     

Extrinsic flexor muscles generate concurrent flexion of all three finger joints

The role of the forearm (extrinsic) finger flexor muscles in initiating rotation of the metacarpophalangeal (MCP) joint and in coordinating flexion at the MCP, the proximal interphalangeal (PIP), and distal interphalangeal (DIP) joints remains a matter of some debate. To address the biomechanical feasibility of the extrinsic flexors performing these actions, a computer simulation of the index finger was created. The model consisted of a planar open-link chain comprised of three revolute joints and four links, driven by the change in length of the flexor muscles. Passive joint characteristics, included in the model, were obtained from system identification experiments involving the application of angular perturbations to the joint of interest. Simulation results reveal that in the absence of passive joint torque, shortening of the extrinsic flexors results in PIP flexion (80°), but DIP (8°) and MCP (7°) joint extension. The inclusion of normal physiological levels of passive joint torque, however, results in simultaneous flexion of all three joints (63° for DIP, 75° for PIP, and 43° for MCP). Applicability of the simulation results was confirmed by recording finger motion produced by electrical stimulation of the extrinsic flexor muscles for the index finger. These findings support the view that the extrinsic flexor muscles can initiate MCP flexion, and produce simultaneous motion at the MCP, PIP, and DIP joints.     

Kinematic evaluation of the finger’s interphalangeal joints coupling mechanism—variability,flexion–extension differences,triggers, locking swanneck deformities,anthropometric correlations

The human finger contains tendon/ligament mechanisms essential for proper control. One mechanism couples the movements of the interphalangeal joints when the (unloaded) finger is flexed with active deep flexor. This study’s aim was to accurately determine in a large finger sample the kinematics and variability of the coupled interphalangeal joint motions, for potential clinical and finger model validation applications. The data could also be applied to humanoid robotic hands. Sixty-eight fingers were measured in seventeen hands in nine subjects. Fingers exhibited great joint mobility variability, with passive proximal interphalangeal hyperextension ranging from zero to almost fifty degrees. Increased measurement accuracy was obtained by using marker frames to amplify finger segment motions. Gravitational forces on the marker frames were not found to invalidate measurements. The recorded interphalangeal joint trajectories were highly consistent, demonstrating the underlying coupling mechanism. The increased accuracy and large sample size allowed for evaluation of detailed trajectory variability, systematic differences between flexion and extension trajectories, and three trigger types, distinct from flexor tendon triggers, involving initial flexion deficits in either proximal or distal interphalangeal joint. The experimental methods, data and analysis should advance insight into normal and pathological finger biomechanics (e.g., swanneck deformities), and could help improve clinical differential diagnostics of trigger finger causes. The marker frame measuring method may be useful to quantify interphalangeal joints trajectories in surgical/rehabilitative outcome studies. The data as a whole provide the most comprehensive collection of interphalangeal joint trajectories for clinical reference and model validation known to us to date.     

Finger joint force minimization in pianists using optimization techniques

A numerical optimization procedure was used to determine finger positions that minimize and maximize finger tendon and joint force objective functions during piano play. A biomechanical finger model for sagittal plane motion, based on finger anatomy, was used to investigate finger tendon tensions and joint reaction forces for finger positions used in playing the piano. For commonly used piano key strike positions, flexor and intrinsic muscle tendon tensions ranged from 0.7 to 3.2 times the fingertip key strike force, while resultant inter-joint compressive forces ranged from 2 to 7 times the magnitude of the fingertip force. In general, use of a curved finger position, with a large metacarpophalangeal joint flexion angle and a small proximal interphalangeal joint flexion angle, reduces flexor tendon tension and resultant finger joint force.     

Finger joint coordination during tapping

We investigated finger joint coordination during tapping by characterizing joint kinematics and torques in terms of muscle activation patterns and energy profiles. Six subjects tapped with their index finger on a computer keyswitch as if they were typing on the middle row of a keyboard. Fingertip force, keyswitch position, kinematics of the metacarpophalangeal (MCP) and the proximal and distal interphalangeal (IP) joints, and intramuscular electromyography of intrinsic and extrinsic finger muscles were measured simultaneously. Finger joint torques were calculated based on a closed-form Newton–Euler inverse dynamic model of the finger. During the keystroke, the MCP joint flexed and the IP joints extended before and throughout the loading phase of the contact period, creating a closing reciprocal motion of the finger joints. As the finger lifted, the MCP joint extended and the interphalangeal (IP) joints flexed, creating an opening reciprocal motion. Intrinsic finger muscle and extrinsic flexor activities both began after the initiation of the downward finger movement. The intrinsic finger muscle activity preceded both the IP joint extension and the onset of extrinsic muscle activity. Only extrinsic extensor activity was present as the finger was lifted. While both potential energy and kinetic energy are present and large enough to overcome the work necessary to press the keyswitch, the motor control strategies utilize the muscle forces and joint torques to ensure a successful keystroke.     

Effects of movement speed and joint position on knee flexor torque in healthy and post-surgical subjects

Coactivation of knee flexors during knee extension assists in joint stability by exerting an opposing torque to the anterior tibial displacement induced by the quadriceps. This opposing torque is believed to be generated by eccentric muscle actions that stiffen the knee, thereby attenuating strain to joint ligaments, particularly the anterior cruciate ligament (ACL). However, as the lengths of knee muscles vary with changes in joint position, the magnitude of flexor/extensor muscle force coupling may likewise vary, possibly affecting the capacity for active knee stabilization. The purpose of this study was to assess the effect of changes in movement speed and joint position on eccentric/concentric muscle action relationships in the knees of uninjured (UNI) and post-ACL-surgery (INJ) subjects (n = 14). All subjects were tested for maximum eccentric and concentric torque of the contralateral knee flexors and extensor muscles at four isokinetic speeds (15 degrees-60 degrees x s(-1)) and four joint position intervals (20 degrees-60 degrees of knee flexion). Eccentric flexor torque was normalized to the percentage of concentric flexor torque generated at each joint position interval for each speed tested (flexor E-C ratio). In order to estimate the capacity of the knee flexors to resist active knee extension, the eccentric-flexor/concentric-extensor ratios were also computed for each joint position interval and speed (flexor/extensor E-C ratio). The results revealed that eccentric torque surpassed concentric torque by 3%-144% across movement speeds and joint position intervals. The magnitude of the flexor E-C ratio and flexor/extensor E-C increased significantly with speed in both groups of subjects (P < 0.05) and tended to rise with muscle length as the knee was extended; peak values were generated at the most extended joint position (20 degrees-30 degrees). Although torque development patterns were symmetrical between the contralateral limbs in both groups, between-group comparisons revealed significantly higher flexor/extensor E-C ratios for the INJ group compared to the UNI group (P < 0.05), particularly at the fastest speed tested (60 degrees x s(-1)). The results indicate that joint position and movement speed influence the eccentric/concentric relationships of knee flexors and extensors. The INJ subjects appeared to accommodate to surgery by developing the eccentric function of their ACL and normal knee flexors, particularly at higher speeds and at more extended knee joint positions. This may assist in the dynamic stabilization of the knee at positions where ACL grafts have been reported to be most vulnerable to strain.     

A combined regimen of controlled motion following flexor tendon repair in "no man's land"

A program of controlled motion following repair of flexor tendons in the hand is presented. This regimen incorporates the features of active extension against rubber band passive flexion, as well as those of controlled passive extension and passive flexion. In this prospective study, 44 digits with complete lacerations of the flexor digitorum profundus and flexor digitorum superficialis in zone 2 were treated. Using the Strickland formula of total active motion of the interphalangeal joints, 36 fingers (82 percent) were rated "excellent"; 7 fingers (16 percent) were rated "good"; 1 finger (2 percent) was rated "fair"; none was rated "poor". There was no statistical difference between the results of delayed primary repair and immediate primary repair.     

The effect of finger extensor mechanism on the flexor force during isometric tasks.

The role of the intrinsic finger flexor muscles was investigated during finger flexion tasks. A suspension system was used to measure isometric finger forces when the point of force application varied along fingers in a distal-proximal direction. Two biomechanical models, with consideration of extensor mechanism Extensor Mechanism Model (EMM) and without consideration of extensor mechanism Flexor Model (FM), were used to calculate forces of extrinsic and intrinsic finger flexors. When the point of force application was at the distal phalanx, the extrinsic flexor muscles flexor digitorum profundus, FDP, and flexor digitorum superficialis, FDS, accounted for over 80% of the summed force of all flexors, and therefore were the major contributors to the joint flexion at the distal interphalangeal (DIP), proximal interphalangeal (PIP), and metacarpophalangeal (MCP) joints. When the point of force application was at the DIP joint, the FDS accounted for more than 70% of the total force of all flexors, and was the major contributor to the PIP and MCP joint flexion. When the force of application was at the PIP joint, the intrinsic muscle group was the major contributor for MCP flexion, accounting for more than 70% of the combined force of all flexors. The results suggest that the effects of the extensor mechanism on the flexors are relatively small when the location of force application is distal to the PIP joint. When the external force is applied proximally to the PIP joint, the extensor mechanism has large influence on force production of all flexors. The current study provides an experimental protocol and biomechanical models that allow estimation of the effects of extensor mechanism on both the extrinsic and intrinsic flexors in various loading conditions, as well as differentiating the contribution of the intrinsic and extrinsic finger flexors during isometric flexion.     

The effect of tendon loading on in-vitro carpal kinematics of the wrist joint

Measurements of in-vitro carpal kinematics of the wrist provide valuable biomechanical data. Tendon loading is often applied during cadaver experiments to simulate natural stabilizing joint compression in the wrist joint. The purpose of this study was to investigate the effect of tendon loading on carpal kinematics in-vitro.A cyclic movement was imposed on 7 cadaveric forearms while the carpal kinematics were acquired by a 4-dimensional rotational X-ray imaging system. The extensor- and flexor tendons were loaded with constant force springs of 50 N, respectively. The measurements were repeated without a load on the tendons. The effect of loading on the kinematics was tested statistically by using a linear mixed model.During flexion and extension, the proximal carpal bones were more extended with tendon loading. The lunate was on the average 2.0° (p=0.012) more extended. With tendon loading the distal carpal bones were more ulnary deviated at each angle of wrist motion. The capitate was on the average 2.4° (p=0.004) more ulnary deviated.During radioulnar deviation, the proximal carpal bones were more radially deviated with the lunate 0.7° more into radial deviation with tendon loading (p<0.001). Conversely, the bones of distal row were more flexed and supinated with the capitate 1.5° more into flexion (p=0.025) and 1.0° more into supination (p=0.011).In conclusion, the application of a constant load onto the flexor and extensor tendons in cadaver experiments has a small but statistically significant effect on the carpal kinematics during flexion–extension and radioulnar deviation.     

Kinetics of crimp and slope grip in rock climbing

The aim was to investigate differences of the kinetics of the crimp and the slope grip used in rock climbing. Nine cadaver fingers were prepared and fixated with the proximal phalanx in a frame. The superficial (FDS) and deep (FDP) flexor tendons were loaded selectively and together with 40 N in the crimp grip (PIP joint flexed 90°/DIP joint hyperextended) and the slope grip position (<25° flexed/50° flexed respectively). Five different grip sizes were tested and the flexion force which was generated to the grip was measured. In the crimp grip the FDP generated more flexion force in small sized holds whereas the FDS generated more force in the larger holds. During the slope grip the FDP was more effective than the FDS. While both tendons were loaded, the flexion force was always greater during crimp grip compared with the slope grip. The FDP seems to be most important for very small holds using the crimp grip but also during slope grip holds whereas the FDS is more important for larger flat holds.     

A dynamic model for finger interphalangeal coordination

In this paper a dynamic model to investigate interphalangeal coordination in the human finger is proposed. Suitable models which describe the relationship between the tendon displacement and the joint angles have been chosen and incorporated into the skeletal dynamic model. A kinematic and kinetic model for interphalangeal coordination is suggested. Digital computer simulations are carried out to study interphalangeal (IP) flexion. Moreover, the effect of two different optimization methods is contrasted. The two optimization algorithms are employed to obtain a set of feasible values for the forces in the tendons or muscles of the finger.     

Modelling tendon excursions and moment arms of the finger flexors: anatomic fidelity versus function

A detailed musculoskeletal model of the human hand is needed to investigate the pathomechanics of tendon disorders and carpal tunnel syndrome. The purpose of this study was to develop a biomechanical model with realistic flexor tendon excursions and moment arms. An existing upper extremity model served as a starting point, which included programmed movement of the index finger. Movement capabilities were added for the other fingers. Metacarpophalangeal articulations were modelled as universal joints to simulate flexion/extension and abduction/adduction while interphalangeal articulations used hinges to represent flexion. Flexor tendon paths were modelled using two approaches. The first method constrained tendons with control points, representing annular pulleys. The second technique used wrap objects at the joints as tendon constraints. Both control point and joint wrap models were iteratively adjusted to coincide with tendon excursions and moment arms from a anthropometric regression model using inputs for a 50th percentile male. Tendon excursions from the joint wrap method best matched the regression model even though anatomic features of the tendon paths were not preserved (absolute differences: mean<0.33 mm, peak<0.74 mm). The joint wrap model also produced similar moment arms to the regression (absolute differences: mean<0.63 mm, peak<1.58 mm). When a scaling algorithm was used to test anthropometrics, the scaled joint wrap models better matched the regression than the scaled control point models. Detailed patient-specific anatomical data will improve model outcomes for clinical use; however, population studies may benefit from simplified geometry, especially with anthropometric scaling.     

Tendon displacements during voluntary and involuntary finger movements

In the human hand, independent movement control of individual fingers is limited. One potential cause for this is mechanical connections between the tendons and muscle bellies corresponding to the different fingers. The aim of this study was to determine the tendon displacement of the flexor digitorum superficialis (FDS) of both the instructed and the neighboring, non-instructed fingers during single finger flexion movements. In nine healthy subjects (age 22–29 years), instructed and non-instructed FDS finger tendon displacement of the index, middle and ring finger was measured using 2D ultrasound analyzed with speckle tracking software in two conditions: active flexion of all finger joints with all fingers free to move and active flexion while the non-instructed fingers were restricted. Our results of the free movement protocol showed an average tendon displacement of 27 mm for index finger flexion, 21 mm for middle finger flexion and 17 mm for ring finger flexion. Displacements of the non-instructed finger tendons (≈12 mm) were higher than expected based of the amount of non-instructed finger movement. In the restricted protocol, we found that, despite minimal joint movements, substantial non-instructed finger tendon displacement (≈9 mm) was still observed, which was interpreted as a result of tendon strain. When this strain component was subtracted from the tendon displacement of the non-instructed fingers during the free movement condition, the relationship between finger movement and tendon displacement of the instructed and non-instructed finger became comparable. Thus, when studying non-instructed finger tendon displacement it is important to take tendon strain into consideration.     

Modality-specific organization for cutaneous and proprioceptive sense in human primary sensory cortex studied by chronic epicortical recording

Modality specificity of human primary somatosensory cortex was studied by recording somatosensory evoked potentials (SEPs) from subdural electrodes in a patient with intractable focal motor seizure. A newly developed device was used for selectively activating proprioception. The spatial and temporal distributions of proprioception-related SEPs elicited by brisk passive flexion movement at the proximal interphalangeal (PIP) joint of the middle finger (4 degrees in 25 ms) were quite different from those to cutaneous sense evoked by electric stimulation of the digital nerve at the same site. It was for the first time demonstrated that proprioception-related SEPs following passive finger movement do not originate in area 3b, which was clearly activated by cutaneous stimulation, and that other sites at the sensorimotor cortex such as areas 2, 3a and 4 possibly contribute to the cortical processing of proprioception.     

Further experience in rehabilitation of zone II flexor tendon repair with dynamic traction splinting

A review of all flexor tendon repairs in the "no man's land" performed from January of 1985 to June of 1987 was done to evaluate the efficacy of our method of rehabilitation. There were 60 fingers (57 patients) with complete laceration of the flexor digitorum profundus and flexor digitorum superficialis tendons in zone II. Fingers with phalangeal fractures, joint injuries, or significant skin loss were excluded. Follow-up ranged from 12 to 48 months. Rehabilitation consisted of a 12-week protocol using the U.S. military combined regimen of controlled motion. Features from the technique of controlled active extension against rubber band passive flexion as well as those of controlled passive extension and passive flexion were incorporated. The palmar pulley modification of Kleinert's dynamic traction splint was utilized. Strickland's total active motion formula was employed to determine results. The results were classified into the four categories of excellent, good, fair, and poor. Fifty-two fingers (86 percent) were rated excellent, 4 fingers (7 percent) were rated good, 1 finger (2 percent) was rated fair, and 3 fingers (5 percent) were rated poor.     

Quantification of fingertip force reduction in the forefinger following simulated paralysis of extensor and intrinsic muscles

Objective estimates of fingertip force reduction following peripheral nerve injuries would assist clinicians in setting realistic expectations for rehabilitating strength of grasp. We quantified the reduction in fingertip force that can be biomechanically attributed to paralysis of the groups of muscles associated with low radial and ulnar palsies. We mounted 11 fresh cadaveric hands (5 right, 6 left) on a frame, placed their forefingers in a functional posture (neutral abduction, 45° of flexion at the metacarpophalangeal and proximal interphalangeal joints, and 10° at the distal interphalangeal joint) and pinned the distal phalanx to a six-axis dynamometer. We pulled on individual tendons with tensions up to 25% of maximal isometric force of their associated muscle and measured fingertip force and torque output. Based on these measurements, we predicted the optimal combination of tendon tensions that maximized palmar force (analogous to tip pinch force, directed perpendicularly from the midpoint of the distal phalanx, in the plane of finger flexion–extension) for three cases: non-paretic (all muscles of forefinger available), low radial palsy (extrinsic extensor muscles unavailable) and low ulnar palsy (intrinsic muscles unavailable). We then applied these combinations of tension to the cadaveric tendons and measured fingertip output. Measured palmar forces were within 2% and 5° of the predicted magnitude and direction, respectively, suggesting tendon tensions superimpose linearly in spite of the complexity of the extensor mechanism. Maximal palmar forces for ulnar and radial palsies were 43 and 85% of non-paretic magnitude, respectively (p<0.05). Thus, the reduction in tip pinch strength seen clinically in low radial palsy may be partly due to loss of the biomechanical contribution of forefinger extrinsic extensor muscles to palmar force. Fingertip forces in low ulnar palsy were 9° further from the desired palmar direction than the non-paretic or low radial palsy cases (p<0.05).     

Effects of knee joint angle on the fascicle behavior of the gastrocnemius muscle during eccentric plantar flexions

The present study aimed to clarify the effects of knee joint angle on the behavior of the medial gastrocnemius muscle (MG) fascicles during eccentric plantar flexions. Eight male subjects performed maximal eccentric plantar flexions at two knee positions [fully extended (K0) and 90° flexed (K90)]. The eccentric actions were preceded by static plantar flexion at a 30° plantar flexed position and then the ankle joint was forcibly dorsiflexed to 15° of dorsiflexion with an isokinetic dynamometer at 30°/s and 150°/s. Tendon force was calculated by dividing the plantar flexion torque by the estimated moment arm of the Achilles tendon. The MG fascicle length was determined with ultrasonography. The tendon forces during eccentric plantar flexions were influenced by the knee joint angle, but not by the angular velocity. The MG fascicle lengths were elongated as the ankle was dorsiflexed in K0, but in K90 they were almost constant despite the identical range of ankle joint motion. These results suggested that MG fascicle behavior during eccentric actions was markedly affected by the knee joint angle. The difference in the fascicle behavior between K0 and K90 could be attributed to the non-linear force–length relations and/or to the slackness of tendinous tissues.     

Estimation of finger muscle tendon tensions and pulley forces during specific sport-climbing grip techniques

The present work displayed the first quantitative data of forces acting on tendons and pulleys during specific sport-climbing grip techniques. A three-dimensional static biomechanical model was used to estimate finger muscle tendon and pulley forces during the "slope" and the "crimp" grip. In the slope grip the finger joints are flexed, and in the crimp grip the distal interphalangeal (DIP) joint is hyperextended while the other joints are flexed. The tendons of the flexor digitorum profundus and superficialis (FDP and FDS), the extensor digitorum communis (EDC), the ulnar and radial interosseus (UI and RI), the lumbrical muscle (LU) and two annular pulleys (A2 and A4) were considered in the model. For the crimp grip in equilibrium conditions, a passive moment for the DIP joint was taken into account in the biomechanical model. This moment was quantified by relating the FDP intramuscular electromyogram (EMG) to the DIP joint external moment. Its intensity was estimated at a quarter of the external moment. The involvement of this parameter in the moment equilibrium equation for the DIP joint is thus essential. The FDP-to-FDS tendon-force ratio was 1.75:1 in the crimp grip and 0.88:1 in the slope grip. This result showed that the FDP was the prime finger flexor in the crimp grip, whereas the tendon tensions were equally distributed between the FDP and FDS tendons in the slope grip. The forces acting on the pulleys were 36 times lower for A2 in the slope grip than in the crimp grip, while the forces acting on A4 were 4 times lower. This current work provides both an experimental procedure and a biomechanical model that allows estimation of tendon tensions and pulley forces crucial for the knowledge about finger injuries in sport climbing.     

Wrist and digital joint motion produce unique flexor tendon force and excursion in the canine forelimb

The force and excursion within the canine digital flexor tendons were measured during passive joint manipulations that simulate those used during rehabilitation after flexor tendon repair and during active muscle contraction, simulating the active rehabilitation protocol. Tendon force was measured using a small buckle placed upon the tendon while excursion was measured using a suture marker and video analysis method. Passive finger motion imposed with the wrist flexed resulted in dramatically lower tendon force (approximately 5 N) compared to passive motion imposed with the wrist extended (approximately 17 N). Lower excursions were seen at the level of the proximal interphalangeal joint with the wrist flexed (approximately 1.5 mm) while high excursion was observed when the wrist was extended or when synergistic finger and wrist motion were imposed (approximately 3.5 mm). Bivariate discriminant analysis of both force and excursion data revealed a natural clustering of the data into three general mechanical paradigms. With the wrist extended and with either one finger or four fingers manipulated, tendons experienced high loads of approximately 1500 g and high excursions of approximately 3.5 mm. In contrast, the same manipulations performed with the wrist flexed resulted in low tendon forces (4-8 N) and low tendon excursions of approximately 1.5 mm. Synergistic wrist and finger manipulation provided the third paradigm where tendon force was relatively low (approximately 4 N) but excursion was as high as those seen in the groups which were manipulated with the wrist extended. Active muscle contraction produced a modest tendon excursion (approximately 1 mm) and high or low tendon force with the wrist extended or flexed, respectively. These data provide the basis for experimentally testable hypotheses with regard to the factors that most significantly affect functional recovery after digital flexor tendon injury and define the normal mechanical operating characteristics of these tendons.     

Finger joint impedance during tapping on a computer keyswitch

We studied the dynamic behavior of finger joints during the contact period of tapping on a computer keyswitch, to characterize and parameterize joint function with a lumped-parameter impedance model. We tested the hypothesis that the metacarpophalangeal (MCP) and interphalangeal (IP) joints act similarly in terms of kinematics, torque, and energy production when tapping. Fifteen human subjects tapped with the index finger of the right hand on a computer keyswitch mounted on a two-axis force sensor, which measured forces in the vertical and sagittal planes. Miniature fiber-optic goniometers mounted across the dorsal side of each joint measured joint kinematics. Joint torques were calculated from endpoint forces and joint kinematics using an inverse dynamic algorithm. For each joint, a linear spring and damper model was fitted to joint torque, position, and velocity during the contact period of each tap (22 per subject on average). The spring-damper model could account for over 90% of the variance in torque when loading and unloading portions of the contact were separated, with model parameters comparable to those previously measured during isometric loading of the finger. The finger joints functioned differently, as illustrated by energy production during the contact period. During the loading phase of contact the MCP joint flexed and produced energy, whereas the proximal and distal IP joints extended and absorbed energy. These results suggest that the MCP joint does work on the interphalangeal joints as well as on the keyswitch.